Austrobaileya

Volume 8

Number 3 2011

AUSTROBAIIEYA

A Journal of Plant Systematics

and

Conservation Biology

Queensland Herbarium

Department of Environment and Resource Management

Queensland

Government

Volume 8

Number 3 2011

A Journal of Plant Systematics

and

Conservation Biology

Queensland Herbarium

Department of Environment and Resource Management

Queensland

Government

Editorial Committee

P.I.Forster (editor)
P.D.Bostock (technical advisor)
G.P.Guymer (technical advisor)

Graphic Design

Will Smith

Desktop Publishing

Yvonne Smith

Austrobaileya

Vol. 1, No. 1 was published on 1 December 1977
Vol. 8, No. 2 was published on 7 December 2010
Vol. 8, No. 3 was published on c. 5 December 2011
Austrobaileya is published once per year.

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addressed to: The Editor, Austrobaileya, Queensland Herbarium, Department of Environment
& Resource Management, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland
4066, Australia. Email: paul.forster@derm.qld.gov.au

ISSN 0155-4131

© Queensland Herbarium 2011

Web site: www.derm.qld.gov.au/herbarium

Austrobaileya is the journal of the Queensland Herbarium and publishes peer-reviewed research
on plants, algae, fungi and lichens (systematics, morphology, geography, anatomy, karyology,
conservation biology and botanical history), with special emphasis on taxa from Queensland.

Opinions expressed by authors are their own and do not necessarily represent the policies or
view of the Queensland Herbarium, Department of Environment & Resource Management.

Austrobaileya 8 ( 3 ): 225-440 ( 2011 )

Contents

A taxonomic revision of Symplocos Jacq. (Symplocaceae) in Australia

L.W. Jessup . 225-251

A taxonomic revision of Pseudoweinmannia Engl. (Cunoniaceae: Geissoieae )

A.C.Rozefelds & B.Pellow . 252-266

A taxonomic revision of Nyssanthes R.Br. (Amaranthaceae)

A. R. Be an & C. II. Miller . 267-279

A taxonomic revision of Pterocaulon section Monenteles (Labill.) Kuntze
(Asteraceae: Inuleae-Plucheinae)

A. R. Bean . 280-334

Heliotropium microspermum E.J.Thomps. (Boraginaceae), a new species
from Queensland

E. J.Thompson . 335-339

Two new species of Pluchea Cass. (Asteraceae: Plucheinae) from
Queensland, Australia

A. R. Bean . 340-346

Eucalyptus melanophloia subsp. nana D.Nicolle & Kleinig, a new mallee
ironbark (E. series Siderophloiae ; Myrtaceae) from central Australia and
north western Queensland

D.Nicolle & D.A.Kleinig . 347-355

Cycas terryana P.I.Forst. (Cycadaceae), a new species from central Queensland

P.I.Forster . 356-363

A taxonomic revision of Trichosanthes L. (Cucurbitaceae) in Australia,
including one new species from the Northern Territory

W.E.Cooper & H.J.de Boer . 364-386

Five new species of Plectranthus L.Her. (Lamiaceae) from New South Wales
and Queensland

P.I.Forster . 387-404

Wilkiea kaarruana Zich & A.J.Ford (Monimiaceae), a new species from
north-east Queensland

F. Zich & A.J.Ford . 405-411

New and reinstated species of the Solanum ellipticum R.Br. (Solanaceae)
species group

A. R. Be an .412-430

Luvunga monophylla (DC.) Mabb. (Rutaceae): a new species for
Queensland

D.G.Fell & D.J.Stanton . 431-434

Hydrocharis dubia (Blume) Backer (Hydrocharitaceae) is an alien species
in Australia

A. R. Be an . 435-437

Reinstatement of Vigna suberecta Benth. (Fabaceae: Phaseoleae)

A. E. Holland . 438-440

A taxonomic revision of Symplocos Jacq.
(Symplocaceae) in Australia

L.W. Jessup

Summary

Jessup, L.W. (2011). A taxonomic revision of Symplocos Jacq. (Symplocaceae) in Australia.
Austrobaileya 8(3): 225-251. The Australian taxa in the genus Symplocos are revised. Nineteen
species are recognised and separate identification keys are provided for flowering and fruiting material.
Symplocos boonjee Jessup, S. bullata Jessup and S. oresbia Jessup are described as new species, four
taxa are raised in rank to species viz. S. gittinsii Jessup, S. glabra Jessup, S. puberula Jessup and
S. wooroonooran Jessup and one new variety, S. cyanocarpa var. pilosa Jessup is described.

Key Words: Symplocaceae, Symplocos , Australia flora, Queensland flora. New South Wales flora,
Symplocos boonjee, Symplocos bullata, Symplocos cyanocarpa var. pilosa, Symplocos gittinsii,
Symplocos glabra, Symplocos oresbia, Symplocos puberula, Symplocos wooroonooran, new species,
identification key

Laurence W. Jessup, Queensland Herbarium, Department of Environment and Resource Management,
Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email:
Laurence.Jessup@derm.qld.gov.au

Introduction

Symplocos Jacq. is a predominantly
pantropical genus of over 320 species with
highest species diversity in Asia, Malesia and
the New World (Wang et al. 2004; Fritsch et
al. 2006, 2008). Symplocos has been usually
regarded as the sole generic representative
of the Symplococeae with division into four
subgenera and various sections based upon
floral structure (Brand 1901). Molecular
phylogenies to date have not supported
this classification and have resulted in the
elevation of Symplocos section Cordyloblaste
to generic rank (as Cordyloblaste Hensch.
ex Moritzi) and the recognition of only two
subgenera and relatively few sections and
series in Symplocos (Fritsch et al. 2008). This
infrageneric classification of the genus is
based on DNA sequence data from five genic
regions (for 90 species) and 51 morphological
characters (Fritsch et al. 2008). For the
Australian region, all taxa can be classified
into Symplocos (subgenus Symplocos)
section Lodhra G.Don (Fritsch et al. 2008)
and represents a major departure from that
proposed by Nooteboom (1975).

Accepted for publication 2 September 2011

Mueller (1862) was the first to recognise
Symplocos in Australia ( S. thwaitesii F.Muell.)
and followed this with a second new species (S.
stawellii F.Muell.) (Mueller 1865). Bentham
(1869) recognised Symplocos thwaitesii but
applied the name S. spicata var. australis
Benth. to specimens of S. stawellii. Bailey
(1890) described Symplocos paucistaminea
F.Muell. & F.M.Bailey ex F.M.Bailey and in
Bailey (1900b) treated the other Australian
species in the same sense as Bentham
(1869). Brand (1901) reinstated the name
Symplocos stawellii and described the Lord
Howe Island species S. candelabrum Brand.
Nooteboom (1975) revised the Old World
species creating a large number of synonyms
and four subspecies and thirty-four varieties
under the apparently polymorphic species
Symplocos cochinchinensis (Lour.) S.Moore.
He followed with a modified treatment of
Australian Symplocos in Nooteboom (1981).
Smith (1978, 1982) chose not to follow
Nooteboom’s broad taxonomic concept in his
treatment of the Fijian species as did Green
(1994) in the Oceanic Islands volume of the
Flora of Australia.

Since 1981 new taxa from Australia have
been recognised and some taxa that were
placed in synonymy by Nooteboom have
continued to be recognised as distinct in

226

Austrobaileya 8(3): 225-251 (2011)

the field and in the herbarium particularly
as more collections have been made (Jessup
1993, 1994, 1997, 2002, 2007, 2010). Also
all of the Australian varieties described by
Nooteboom under Symplocos cochinchinensis
subsp. thwaitesii (F.Muell.) Noot. are now
well-collected throughout their natural range
and are represented in the herbarium by
quality specimens that exhibit several distinct
correlating character states sufficient to allow
recognition as species. In this paper four
previously described taxa are raised in rank
to species and three new species and one new
variety are described.

Materials and methods

This paper is based on herbarium collections in
BM, BRI, CANB, CNS (previously QRS), K,
L, MEL and NSW and some field observations
by the author. All specimens cited have been
seen by the author. Descriptions of flowers
were prepared from material preserved
in FAA or 70% alcohol and glycerol or
reconstituted by briefly boiling in water.
Descriptions of fruit were prepared from
both dried and alcohol preserved material.
Common abbreviations in the specimen
citations are F.R. (Forest Reserve), L.A.
(Logging Area), N.R (National Park), S.F.R./
S.F. (State Forest Reserve/State Forest), T.R.
(Timber Reserve). The abbreviation NCA
is for the Queensland Nature Conservation
Act 1992 and its associated regulations. The
abbreviation EPBC refers to the Environment
Protection and Biodiversity Conservation Act
1999

Taxonomy

Symplocos Jacq., Enum. PI. Carib. 5, 24
(1760). Type species: S. martinicensis Jacq.

Trees or shrubs, dioecious, polygamous
or hermaphroditic. Leaves mostly spirally
arranged, simple, penninerved, serrate,
dentate, or entire, exstipulate. Inflorescence
spicate, racemose, a thyrse or a panicle, or
flowers solitary or fasciculate, axillary, rarely
terminal. Flowers actinomorphic, bisexual or
often functionally unisexual, subtended by
a bract and 2 bracteoles. Ovary inferior or
rarely semi-inferior. Hypanthium completely
adnate to ovary. Calyx limb epigynous,
mostly 5-lobed; lobes imbricate, persistent.
Corolla gamopetalous but divided to near
base into 3-10 petals (5 in Australian species),
quincuncially imbricate, white, cream or pink.
Stamens few to many, connate only at base,
adnate to the base of the corolla, pentadelphous
in alternipetalous bundles or (not in Australia)
monadelphous; filaments terete. Anthers
bilocular, introrse, dehiscing longitudinally.
Disc inside staminal ring. Ovary 2-5-locular
(3-locular in Australia) with 2-4 anatropous
ovules per locule. Placentation axile but septa
separating distally. Style 1, stigma simple,
capitate or peltate. Fruit a monopyrenous
drupe, crowned by persistent calyx lobes,
frequently 1-celled and 1-seeded by abortion;
mesocarp fleshy or not; endocarp hard, smooth
or longitudinally ridged, with an apical
depression. Seed with copious endosperm;
embryo straight or curved.

The genus includes 300-350 species in
tropical, subtropical and temperate parts of
Asia, Malesia, North and South America, the
West Indies, the Pacific islands and Australia.
Nineteen species occur in Australia.

Jessup, Symplocos in Australia

227

Fig. 1. Symplocos ampulliformis. A. part of inflorescence with flowers and buds x 4, Gray 3362 (BRI). B. fruit x 2,
Hyland 13459 (BRI). C. leafabaxial surface x fl Volck 1389 (BRI). .S', boonjee. D. inflorescence with flowers x 3, Gray
5575 (CNS). E. flower x 6, Hyland 14345 (CNS). F. fruit x 2, Gray 5575 (CNS). S. bullata. G. inflorescence with flowers
x 3. H. leaf adaxial surface x 1 . 1 . leaf abaxial surface x 1 . G-I from Gray 2543 (BRI). Del. W.Smith.

228

Austrobaileya 8(3): 225-251 (2011)

Key to Australian Symplocos taxa using flowering specimens

1 Inflorescence racemose or a panicle of shortly pedicellate flowers.2

1. Inflorescence spicate or a panicle or fascicle of sessile flowers, or flowers

solitary, axillary.10

2 Lamina margins serrate or dentate, sometimes obscurely denticulate.3

2. Lamina margins denticulate or entire.7

3 Petiole up to 4 mm long.4

3. Petiole 5-17 mm long.5

4 Young branchlets appressed puberulous; petals 1.4—1.6 mm wide;

stamens mostly 15-20.2. S. baeuerlenii

4. Young branchlets sparsely pilose with erect hairs; petals 2-2.5 mm wide;

stamens 30-40 . 11. S. harroldii

5 Young branchlets, petiole and inflorescence glabrous or nearly so.6

5. Young branchlets, petiole and inflorescence appressed puberulous or

pubescent.16. S. puberula

6 Calyx limb 1.7-2 mm long; stamens more than 40; petiole mostly 5-15

mm long.18. S. thwaitesii

6. Calyx limb 0.7-0.8 mm long; stamens less than 40; petiole mostly 20-30

mm long.5. S. candelabrum

7 Bracts up to 1.2 mm long; bracteoles up to 0.6 mm long; petals up to 4 mm

long; stamens up to 20; style up to 2.5 mm long.8

7. Bracts 1-2.5 mm long; bracteoles 0.6-1.2 mm long; petals 4 mm long or

more; stamens 30-50; style 3 mm long or more.9

8 Secondary veins mostly 7-9 pairs; basal ones not more acute and longer

than distal ones; bracts caducous before anthesis; petals 2-2.5 mm long;
disc and style glabrous.

8. Secondary veins mostly 4-6 pairs; basal ones more acute and longer than

distal ones; bracts persistent to after anthesis; petals 3-3.7 mm long;
disc and base of style puberulous.1. S

9 Young branchlets glabrescent; lamina 2-6.5 cm long; inflorescence

appressed puberulous; bracts ovate; style 3-3.5 mm long.19. S. wooroonooran

9. Young branchlets glabrous; lamina 7-15 cm long; inflorescence glabrous;

bracts oblong; style 4.5-5 mm long.8. S. glabra

10 Young branchlets, petiole or inflorescence pilose, villous or pubescent

with erect hairs. 11

10. Young branchlets glabrous or puberulous; petiole glabrous; inflorescence

glabrous, puberulous, or shortly appressed pubescent.16

11 Calyx densely pubescent outside.12

11. Calyx glabrous or sparsely pubescent outside.13

12 Secondary veins 6-12 pairs; inflorescence a few-flowered fascicle

.7. S. cyanocarpa var. pilosa

12. Secondary veins mostly 15-20 pairs; inflorescence spicate with many

flowers.5. S. crassiramifera

. 14. S. oresbia

. ampulliformis

Jessup, Symplocos in Australia 229

13 Sterile shoots rarely with cataphylls; twig hairs, reddish brown; midvein

above pilose or villous.14

13. Sterile shoots bearing cataphylls; twig hairs white, hyaline or pale ginger

brown; midvein glabrous above.15

14 Lamina in mature foliage bullate, margins more or less recurved.4. S. bullata

14. Lamina not bullate, margins flat.15. S. paucistaminea

15 Lamina sometimes bullate between veins, base mostly acutely cuneate,

margins often recurved; bracts c. 2 mm long,; calyx lobes obtuse or

rounded, entire or minutely denticulate.13. S. hylandii

15. Lamina not bullate, base rounded or cordate, margins plane; bracts 3-9

mm long; calyx lobes acute, ciliate.12. S. hayesii

16 Inflorescence axes usually branching above the base;.17

16. Inflorescence axes solitary or clustered, unbranched above the base

or flowers fasciculate or solitary. 21

17 Lamina margins crenulate and serrulate.17. S. stawellii

17. Lamina margins entire, rarely some denticulations evident.18

18 Stamens (12—)15—20.19

18. Stamens 30-70 . 20

19 Secondary veins mostly 7-9 pairs; basal ones not more acute and longer

than distal ones; bracts caducous before anthesis; petals 2-2.5 mm long;

disc and style glabrous.14. S. oresbia

19. Secondary veins mostly 4-6 pairs; basal ones often more acute and

longer than distal ones; bracts persistent to after anthesis; petals 3-3.7

mm long; disc and base of style puberulous.1. S. ampulliformis

20 Petiole 3-10 mm long; inflorescence glabrous; bracts oblong; stamens

mostly 30-50; style 4.5- mm long.9. S. glabra

20. Petiole 10-35 mm long; inflorescence sparsely appressed puberulous;

bracts broadly ovate or depressed ovate; stamens mostly 60-70; style

3.5-4 mm long.8. S. gittinsii

21 Inflorescence usually several clustered axillary spikes to 1.5 cm long;

bracts 1.2-1.5 mm long, sometimes denticulate, pubescent; style 2-2.5

mm long.10. S. graniticola

21. Inflorescence a fascicle, or flowers solitary; bracts 1.5 mm long or

more; style 2.5-3.5 mm long.22

22 Calyx lobes 1-1.4 mm long; petals rounded; filaments pilose

.7. S. cyanocarpa var. cyanocarpa

22. Calyx lobes 4-5.5 mm long; petals acute; filaments glabrous.3. S. boonjee

230

Austrobaileya 8(3): 225-251 (2011)

Key to Australian Symplocos taxa using fruiting specimens

1 Dry fruit ampulliform (flask-shaped).2

1. Dry fruit globose, depressed globose, or ellipsoid to subcylindrical,

rarely ovoid or obovoid.10

2 Young branchlets and petiole pilose or villous; infructescence spicate,

pilose or villous.3

2. Young branchlets and petiole glabrous, puberulous, or appressed
pubescent; infructescence paniculate, or racemose, glabrous or
appressed pubescent.6

3 Dry fruit 5-6 mm long, 3.5-4 mm wide.15. S. paucistaminea

3. Dry fruit 7-10 mm long, 4-5 mm wide.4

4 Secondary veins 15-20 pairs.6. S. crassiramifera

4. Secondary veins 7-12 pairs.5

5 Midvein densely villous on upper (adaxial) surface of lamina.4. S. bullata

5. Midvein glabrous on upper (adaxial) surface of lamina.13. S. hylandii

6 Lamina margins not entire, sometimes teeth scarcely evident on recurved

margins.7

6. Lamina margins entire.8

7 Young branchlets and petiole glabrous; lamina margins flat.18. S. thwaitesii

7. Young branchlets puberulous or pubescent; petiole glabrescent; lamina

margins recurved.16. S. puberula

8 Secondary veins mostly 4-6 pairs; basal pair more acute and longer; style

pubescent at base.1. S. ampulliformis

8. Secondary veins 7-12 pairs, not more acute and longer near base; style

glabrous.9

9 Young branchlets puberulous or pubescent, glabrescent; fruit 5.8-6.3

mm long, 4-5.5 mm wide.19. S. wooroonooran

9. Young branchlets glabrous; fruit 10-15 mm long; 7-9 mm wide.14. S. oresbia

10 Infructescence a glomerule or fruit solitary at each node.11

10. Infructescence spicate or racemose, branched or unbranched.12

11 Fruit glabrous; calyx lobes 4-5.5 mm long.3. S. boonjee

11. Fruit usually not glabrous; calyx lobes 1-1.4 mm long.7. S. cyanocarpa

12 Lamina margins entire or sometimes obscurely denticulate.13

12. Lamina margins serrate or dentate.16

13 Fruit mostly obovoid (Lord Howe Island endemic).5. S. candelabrum

13. Fruit globose, ovoid or ellipsoid (Australian mainland).14

14 Lamina length usually 3-6 times width.10. S. graniticola

14. Lamina length usually less than 3 times width.15

15 Petioles 10-35 mm long. Pedicels 0-0.2 mm long; fruit 6-7 mm long;

4-5 mm wide.

15. Petioles 3-10 mm long. Pedicels 0.2-2 mm long; fruit 9-11 mm long;
8-10 mm wide.

8. S. gittinsii
. 9. S. glabra

231

Jessup, Symplocos in Australia

16 Secondary and tertiary veins raised above (adaxially) (dried leaves).17

16. Secondary and tertiary veins not raised above (adaxially).19

17 Young branchlets, bud scales and infructescence axis glabrous.18. S. thwaitesii

17. Young branchlets, bud scales and infructescence axis not glabrous.18

18 Young branchlets sparsely appressed puberulous; petiole 6-25 mm long;

fruit 3.5-4.5 mm wide.17. S. stawellii

18. Young branchlets sparsely pilose with erect hairs; petiole 3-4 mm long;

fruit 6-7 mm wide.11. S. harroldii

19 Shoots usually without cataphylls; lamina 0.7-2 cm wide, sparsely

appressed puberulous on underside (abaxially), base acutely cuneate;

fruit 4-9 mm long.2. S. baeuerlenii

19. Shoots usually bearing scattered cataphylls; lamina 2.5-5.5 cm wide,

sparsely pilose on underside (abaxially), base rounded or cordate; fruit

15-20 mm long.12. S. hayesii

1. Symplocos ampulliformis C.T.White,
Proc. Roy. Soc. Queensland 50: 81 (1939).
Type: Queensland. Cook District: Mt
Spurgeon, September 1936, C.T.White 10581
(holo: BRI; iso: A n.v .; BM).

Illustrations : Hyland etal. (2010).

Shrub or small tree to 10m. Shoots sparsely
appressed puberulous, glabrescent; young
branchlets glabrous. Leaves chartaceous;
lamina oblanceolate or elliptic, 4.5-12 cm
long, 2-5 cm wide, acuminate at apex,
acute or attenuate at base, glabrous; margins
recurved, entire or nearly so; secondary
veins mostly 4-6 pairs, the basal pair often
longer and more acute; petiole 4-10 mm
long, glabrous. Inflorescence usually a cluster
of racemose axes to 2 cm long, glabrous or
sparsely appressed puberulous, sometimes
shortly branched near base; bracts and
bracteoles ovate, puberulous or ciliate,
persistent to anthesis, bracts c. 0.5 mm long,
bracteoles c. 0.4 mm long. Calyx limb 0.6-0.8
mm long, sparsely puberulous, glabrescent;
lobes depressed ovate or rounded, 0.3-0.5
mm long, glabrescent, margins ciliate. Petals
oblong or narrowly obovate, 2.5-3.5 mm
long, white, glabrous. Stamens (12—)15, 1.5-4
mm long. Disc puberulous. Style c. 2.5 mm
long, puberulous at base. Fruit ampulliform
to narrowly ovoid, 10-15 mm long, 5-6 mm
wide, blue to blue-black or purple-black. Fig.
1A-C.

Additional selected specimens examined: Queensland.
Cook District: N.P.R. 133, summit of Mt Sorrow walking
track. Cape Tribulation, Nov 2000, Ford AF2524 et al.
(BRI); S.F.R.144, Parish of Whypalla, Bower Bird L.A.,
Dec 1987, Hyland 13459 (BRI, CNS); loc. cit., May 1989,
Gray 5046 (BRI, CNS); S.F.R.144, Dasyurid L.A., Apr
1980, Gray 1682 (BRI, CNS); S.F.R.144, Chowchilla
L.A., Jul 1982, Gray 2663 (BRI, CNS); Mt Lewis, SW
of Mossman, Aug 1957, Volck 1389 (BRI); S.F.R.143,
Parish of Riflemead, North Mary L.A., Dec 1984, Gray
3758 (BRI); S.F.R.143, South Mary L.A., Aug 1973,
Irvine 612 (BRI); S.F.R.143, Parish of Riflemead, South
Mary L.A., Gray 3362 (BRI, CNS); S.F.R.143, Parish of
Riflemead, Carbine L.A., Nov 1987, Hyland 25249RFK
(BRI); loc. cit., Jan 1985, Gray 3835 (BRI); S.F.R. 143,
Bushy L.A., Feb 1982, Hyland 11691( BRI); T.R.143,
Zarda L.A., Sep 1973, Hyland 2884RFK (BRI); Along
Falchetti Track to the Mitchell Bomber crash site on
the NE peak of Mt Bartle Frere, Jul 2001, Jago 6011
& Gandini (BRI); Upper E Mt Bartle Frere, Oct 1985,
Godwin C2900 (BRI).

Distribution and habitat: This species is
endemic to Queensland in the Wet Tropics and
occurs on Mt Sorrow near Cape Tribulation,
the Mt Windsor Tableland, Mt Carbine
Tableland, in the vicinity of Mount Lewis,
and on Mt Bartle Frere (Map 1), in upland and
montane notophyll and microphyll vineforest
on granite, granodiorite (all upper montane
localities) and phyllite substrates.

Phenology : Flowers are recorded from March
to May; fruit from January to July.

Notes: This species was treated as Symplocos
cochinchinensis var. montana (C.T.White)
Noot. (as S. cochinchinensis subsp. thwaitesii
var. montana) in Nooteboom (1982).

232

Conservation status : Symplocos ampulliformis
is currently listed as Near Threatened under the

NCA.

Map 1. Distribution of Symplocos ampulliformis

2. Symplocos baeuerlenii R.T.Baker, Proc.
Linn. Soc. New South Wales 27: 594, t. 28
(1902). Type: New South Wales. Tumbulgum,
Murwillumbah, s.dat ., W. Bauerlen s.n. (holo:
NSW; iso: MEL).

Illustrations : Floyd (1989: 396); Leiper et al.
(2008: 336).

Shrub or small tree to 10 m. Shoots and young
branchlets puberulous. Leaves coriaceous;
lamina narrowly elliptic or lanceolate, 1.5-8
cm long, 0.7-2 cm wide, acuminate at apex,
acute at base, glabrous or with a few hairs on
midvein and undersurface; margins mostly
serrate; secondary veins mostly 7-10 pairs;
petiole 2-4 mm long, puberulous, glabrescent.
Inflorescence racemose, to 0.6 cm or flowers
solitary, puberulous; pedicels to 1 mm long;
bracts and bracteoles puberulous, semi-
persistent, bracts acutely triangular, 0.8-1.2
mm long, bracteoles ovate, 0.3-0.5 mm
long. Calyx limb 0.7-1.1 mm long, glabrous;
lobes depressed ovate or rounded, 0.5-0.8
mm long, glabrous. Petals obovate, 2-3 mm
long, 1.4-1.6 mm wide, white, glabrous.
Stamens mostly 15-20, 1.5-3 mm long. Disc
pubescent. Style 1.5-2.5 mm long, pubescent
at base. Fruit ovoid, to 9 mm long, red.

Additional selected specimens examined : Queensland.
Moreton District: Lyrebird Ridge Road, Springbrook
Plateau, Dec 1993, Grimshaw G259 (BRI); Springbrook,
Aug 1945, Blake 15892 (BRI); loc. cit ., Sep 1929, Rudder
4022 (BRI, L); loc. cit., Sep 1929, White 6265 (BRI); loc.

Austrobaileya 8(3): 225-251 (2011)

cit., Aug 1930, White 7059 (BM, BRI); Track to ‘Best of
All’ Lookout, Springbrook, Jul 1977, McDonald 1946 &
Elsol (BRI); End of Currumbin Valley road, Mt Cougal
N.P., Nov 1992, Thomas 96 & Barry (BRI). New South
Wales. Mt Nardi, Goommbar S.F., Feb 1979, Floyd 1194
(BRI); Whian Whian S.F., Jun 1945, White 12758 (BRI);
loc. cit., Jun 1947, Hayes s.n. (BRI [AQ537353]); loc. cit.,
Nov 1949, White s.n. (BRI [AQ335709]); loc. cit., 1953-
1958, Webb & Tracey s.n. (BRI [AQ84368]); Minyon,
near Mullimbimby, Aug 1936, White 10480 (BRI);
Whian Whian, N of Lismore, Jun 1965, Smith 12303 et
al. (BRI); c. 6 km W of Minyon Falls, Whian Whian S.F.,
Jul 1975, Williams s.n. (BRI [AQ504389]).

Distribution and habitat : This species is
endemic to south-east Queensland and north¬
east New South Wales where it occurs from
the Macpherson Range near Springbrook,
south to Lismore (Map 2), in microphyll and
notophyll vineforests on substrates derived
from basalt and enriched rhyolite.

Map 2. Distribution of Symplocos baeuerlenii

Phenology : Flowers are recorded from July
to September; fruit in January and February.

Notes : This species was included under
Symplocos cochinchinensis var. montana
(as S. cochinchinensis subsp. thwaitesii var.
montana ) in Nooteboom (1982).

Conservation status : Symplocos baeuerlenii
is currently listed as Vulnerable under the

NCA.

3. Symplocos boonjee Jessup species nova

S. cyanocarpam simulans sed bracteis
longioribus (2-5 mm contra 1.5-1.7 mm
longis) bracteolis longioribus (2-3 mm contra
0.5-1 mm) sepalis longioribus (4-5.5 mm
contra 1-1.4 mm) petalis ovatis acutis et
filamentis glabris differt. Typus: Queensland.
Cook District: Wooroonooran National Park,
Stockwellia site, 10 July 2000, P.I.Forster

Jessup, Symplocos in Australia

233

PIF25915, R.Booth & R.Jensen (holo: BRI;
isotypi distribuendi).

Symplocos sp. (Boonjie B.P.Hyland 2753);
(Jessup 1994, 1997, 2002, 2007, 2010).

Illustrations : Cooper & Cooper (2004: 538),
under Symplocos sp. (Boonjee); Hyland et
al. (2010), under Symplocos sp. (Boonjie
B.P.Hyland 2753).

Shrub to 3 (-5) m. Shoots, cataphylls and
young branchlets glabrous; cataphylls usually
caducous. Leaves chartaceous; lamina
narrowly obovate or narrowly oblanceolate,
4.5-11 cm long, 1-3 cm wide, acuminate at
apex, acute or attenuate at base, glabrous;
margins usually entire, slightly recurved;
secondary veins mostly 5-7 pairs; petiole

3- 7 mm long, glabrous. Inflorescence a
solitary flower in axil of a leaf or cataphyll;
the axis c. 1.5 mm long with up to 9 bracts
or a fascicle with one open flower surrounded
by several buds; lower bracts broadly ovate to
depressed ovate, upper ones ovate to narrowly
ovate, 3.5-5 mm long, persistent, sparsely
appressed pubescent to nearly glabrous;
margins denticulate, ciliate; bracteoles 2-3
mm long, similar to upper bracts. Calyx limb

4- 5.5 mm long; lobes narrowly ovate, acute,
sparsely puberulous; margin denticulate,
ciliate; tip incurved. Petals narrowly ovate,
acute or obtuse, 3-3.5 mm long, 1.5-1.7 mm
wide, cream to white, glabrous. Stamens
35-40 in males, 25-30 in females, 0.8-3 mm
long, filaments glabrous. Disc pilose. Style
3-3.5 mm long (c. 0.5 mm in males), with a
few hairs at base, otherwise glabrous. Fruit
oblong or ellipsoidal, c. 17 mm long and 3.5
mm wide, glabrous, blue. Fig. 1D-F.

Additional selected specimens examined : Queensland.
Cook District: E side of Mulgrave River, Goldsborough
Valley, Jun 1995, Hunter JH3704 (BRI); Gadgarra S.F.,
S of Butchers Creek, Jun 1995, Hunter JH3904 (BRI);
S.F.R. 310, Swipers F.A., E of Malanda, Aug 1963,
Hyland AFO/2753 (BRI); Swipers Flat, Wooroonooran
N.P., Jul 1995, Forster PIF17138 & Figg (BRI); Boonjee
F.A., near Bartle Frere track, Nov 1988, Jessup GJM362
et al. (BRI); Russell River, Johnson s.n. (MEF46567);
Palmerston Track ridge, Feb 1995 Hunter JH1158 (BRI);
Breeden Property, Hughes Road, Topaz, Mar 1992,
Cooper & Cooper 221 (CNS); T.R. 1230, Bartle Frere,
Boonjee F.A., Sep 1992, Gray 5575 (CNS); S.F.R. 755,
Bartle Frere, Gosschalk F.A., Nov 1991, Hyland 14345
(CNS); R.926, Kauri Reserve, Dirran, Oct 1987, Dansie
20166 (CNS); N.P.R. 904, along track between Coolamon
Creek and Combo Creek, Oct 2000, FordAF2462 (BRI);

Wooroonooran N.P, Combo Creek, Nov 1998, Forster
PIF23990 et al. (BRI).

Distribution and habitat : This species is
endemic to the Wet Tropics of Queensland
where it is known from notophyll and
mesophyll vineforests mostly on the western
side of Wooroonooran N.P, and nearby forest
remnants (Map 3), on metamorphic or basalt
substrates.

Map 3. Distribution of Symplocos boonjee

Phenology: Flowers are recorded from June
to February; fruit in July and August.

Notes: Specimens of this species (e.g. Hyland
2753 ) were tentatively included in Symplocos
cyanocarpa C.T.White in Nooteboom (1982).

Etymology: The species epithet

commemorates a botanically rich but loosely
defined locality on the eastern edge of the
Atherton Tableland known as Boonjee or
Boonjie. For the sake of consistency the first
alternative spelling is used as it has been used
in the names Syzygium boonjee B.Hyland and
Bulbophyllum boonjee B.Gray & D.L. Jones (=
Adelopetalum boonjee (B.Gray & D.L.Jones)
D.L.Jones & M.A.Clem.).

Conservation status: Symplocos boonjee is
only known from Wooroonooran N.P. At a
few sites it has been recorded as common.

234

Additional collections and observations are
required to determine its area of occupancy
and population numbers.

4. Symplocos bullata Jessup species nova

5. paucistamineam simulans sed foliis
bullatis, marginibus recurvis, stylis longibus
(2-2.5 mm, contra 1.2-1.8 mm) et fructibus
majoribus (8-9 x 3.5-4 mm, contra 5-5.8 x
7.5-8 mm) differt. Typus: Queensland. Cook
District: State Forest Reserve 143, North
Mary Logging Area, 25 April 1982, B.Gray
2543 (holo: BRI; iso: BRI, CNS).

Symplocos sp. (North Mary B.Gray 2543);
(Jessup 2002, 2007, 2010).

Illustrations: Hyland et al. (2010), under
Symplocos sp. (North Mary B.Gray 2543).

Dioecious tree to 15 m. Shoots and young
branchlets yellowish brown or reddish-
brown to dark brown villous, cataphylls
rarely evident. Leaves thinly coriaceous
or chartaceous; lamina lanceolate, elliptic,
obovate or oblanceolate, 4-14 cm long, 1.2-5
cm wide, acuminate at apex, cuneate, acute
or scarcely obtusely so or obtuse, truncate
or rounded at base, mature leaves bullate
between main veins, glabrescent above, pilose
below; margins obscurely denticulate or
entire, recurved; secondary veins mostly 7-
12 pairs, usually depressed on upper surface;
petiole 4-11 mm long, pilose. Inflorescences
spicate, to 5 cm, pilose; bracts and bracteoles
ovate, densely reddish-brown pilose or
villous, persistent, bracts 1.8-2.5 mm long,
bracteoles 1.3-1.5 mm long. Calyx limb
1.2-1.5 mm long; lobes ovate, 1-1.2 mm long,
pubescent in middle near apex with reddish-
brown hairs, sometimes glabrescent. Petals
obovate, 3-3.5 mm long, 1—1.5 mm wide,
cream to white, mostly glabrous. Hypanthium
glabrous. Stamens (15-) 20-25 in males, 7-10
in females and apparently sterile, 1-2.5 mm
long. Disc glabrous. Style 2-2.5 mm long,
glabrous; females with peltate stigma. Fruit
broadly ovoid or ampulliform 8-9 mm long,
7.5-8 mm wide, colour unknown. Fig. 1G-I.

Additional selected specimens examined : Queensland.
Cook District: Daintree N.P., NW of Black Mountain,
May 1998, Forster PIF22903 et al. (BRI); Daintree
N.R, near end of Mt Lewis road, Nov 1988, Jessup et al.
GJM155 (BRI); S.F.R.143 Riflemead, North Mary L.A.,

Austrobaileya 8(3): 225-251 (2011)

Jun 1995, Gray 6226 (BRI); loc. c/7., Dec 1995, Gray
6479 (CNS photo); S.F.R. 652, Mt Fisher, May 1975,
Hyland3I75RFK (BRI, CNS).

Distribution and habitat: Endemic to the Wet
Tropics of Queensland where it occurs in the
Mt Lewis to Mt Carbine Tableland area west
of Mossman and with an outlier at Mt Fisher,
south of Millaa Millaa (Map 4). Grows in
montane microphyll and notophyll vineforests
on granite substrates.

Map 4. Distribution of Symplocos bullata

Phenology: Flowers are recorded from April
to June; fruit in December.

Etymology: The specific epithet is derived
from the Latin adjective bullatus (bullate,
blistered or puckered) and refers to the
appearance of the upper surface of the lamina
between the veins.

Conservation status: Symplocos bullata
is only known from the localities cited
above. Additional collections are required
to determine its area of occupancy and
population numbers.

235

Jessup, Symplocos in Australia

5. Symplocos candelabrum Brand in Engl.,
Pflanzenr. 6: 39 (1901); S. cochinchinensis
var. candelabrum (Brand) Noot., Brunonia
4: 323 (1982). Type: New South Wales. Lord
Howe Island, April 1898, J.H.Maiden s.n.
(holo: B [destroyed]; iso: BM, BRI).

Illustration : Green (1994).

Tree to 13 m, usually much smaller. Shoots
and young branchlets glabrous or glabrescent.
Leaves thinly coriaceous; lamina elliptic
or lanceolate, 6-16 cm long, 2-6 cm wide,
acute to acuminate at apex, attenuate at
base, glabrous; margins entire to remotely
serrulate-denticulate near apex, more or less
recurved near base; secondary veins mostly
8-11 pairs; petiole 20-30 mm long, glabrous.
Inflorescence predominately racemose to 6
cm, glabrous or with few very sparse hairs;
pedicels to 5 mm long; bracts and bracteoles
triangular, ciliate; bracts 1—1.5 mm long,
caducous before anthesis; bracteoles 0.8-1.5
mm long, often persisting a little longer
than the bracts. Hypanthium glabrous; calyx
limb 0.6-0.8 mm long; lobes broadly ovate,
glabrous with ciliate margin. Petals obovate,
mostly 6-7 mm long, 3-3.5 mm wide, white,
glabrous. Stamens mostly 30-40, 5-6 mm
long. Disc with a few hairs. Style 4-5 mm
long, pubescent at base. Lruit mostly obovoid,
10-16 mm long, 7-10 mm wide, blue.

Additional selected specimens examined : New South
Wales. Lord Howe Island: Mt Gower walking track,
Nov 2000, Conn 4320 (BRI); Halfway to saddle from
Erskines Creek on summit track, Nov 2000, Le Cussan
1056 (BRI); loc. eit., Feb 2004, Le Cussan 1375 (BRI);
Intermediate Hill track. Mar 2003, Le Cussan 1280
(BRI).

Distribution and habitat: Occurs only in
submontane forest at the southern part of
Lord Howe Island.

Phenology: Llowers are recorded from March
to June; Mature fruit in Lebruary.

Notes: This species was included under
Symplocos cochinchinensis var. candelabrum
(C.T.White)Noot. (as S. cochinchinensis subsp.
thwaitesii var. candelabrum ) in Nooteboom
(1982).

Conservation status: Symplocos candelabrum
is currently not listed under the EPBC Act
despite its highly restricted distribution.

6. Symplocos crassiramifera Noot., Leiden
Bot. Ser. 1: 186 (1975). Type: Queensland.
Cook District: Devil Devil Creek, a few
miles NE of Rumula and c. 8 miles [13.3km]
SSW of Mossman, 1 September 1948, L.S.
Smith 3965 (holo: BRI; iso: L).

Illustrations: Hyland et al. (2010).

Shrub or small tree to 8 m with few branches.
Shoots and young branchlets with caducous
cataphylls, pilose. Leaves coriaceous; lamina
narrowly elliptic to narrowly oblanceolate,
20-41 cm long, 3.5-8.5 cm wide, acuminate at
apex, acute or obtuse at base, glabrous above,
pilose below; margins revolute, remotely
denticulate to entire; secondary veins mostly
15-20 pairs; petiole 8-20 mm long, pilose.
Inflorescences comprising clustered spicate
axes to 15 cm, villous or pilose; bracts and
bracteoles ovate, brown pubescent, persistent,
bracts 2-2.5 mm long, bracteoles 1.7-2
mm long. Calyx limb 1.5-2 mm long, lobes
broadly ovate or rounded, pubescent. Petals
obovate, 4-4.5 mm long, 1.7-2 mm wide,
pink, glabrous. Stamens c. 25, 3-4 mm long.
Disc inconspicuous. Style 2.5-4 mm long,
pubescent at base. Lruit ampulliform, 7-10
mm long, 4-5 mm wide, pink.

Additional selected specimens examined : Queensland.
Cook District: Little Mossman L.A., S.F.143, Jul 1994,
Forster PIF15513 et al. (BRI); Rex Range, c. 1 km
from Mossman-Julatten Road, on forest track c. 9 km
NE of Julatten, Dec 1988, Jessup 880 et al. (BRI); Rex
Highway, near Julatten, Apr 1962, Webb & Tracey 5961
(BRI); The Gate, property of G. & S. Kurmond, 105
Salisbury Drive, Flin Creek, Julatten, Jan 2002, Ford
AF3260 (BRI); T.R. 66, Mt Lewis, May 1979, Moriarty
2603 (BRI, CNS); 7 km along Mt Lewis Road, Mt Lewis
F.R., Apr 2002, Booth 3092 & Jensen (BRI); T.R. 66, 8
km along Mt Lewis Road, Jun 1997, Forster P1F21233 et
al. (BRI); Mt Lewis forestry road, SW of Mossman, May

1972, Wrigley & Telford NQ140 (BRI, CANB); Rumula,
Apr 1962, McKee 9106 (BRI); Portion 203 Garioch, Jun

1973, Hyland 6758 (BRI, CNS).

Distribution and habitat: Endemic to the
Wet Tropics of Queensland where it is known
only from the vicinity of the headwaters of
the South Mossman River to Mt Lewis (Map
5), in notophyll and mesophyll vineforests on
granite and metamorphic substrates.

236

Map 5. Distribution of Symplocos crassiramifera

Phenology : Flowers are recorded from May
to July; fruit in September.

Conservation status : Symplocos crassiramifera
is currently listed as Vulnerable under the NCA.

7. Symplocos cyanocarpa C.T.White, Proc.
Roy. Soc. Queensland 47: 69 (1936). Type:
Queensland. Cook District: Mossman River,
4 February 1932, L.J.Brass 2050 (holo: A,
n.v.\ iso: BRI, MEL).

Shrub or small tree to 5 m. Shoots and young
branchlets pilose, glabrescent or glabrous,
sometimes with pubescent cataphylls.
Leaves chartaceous to coriaceous; lamina

Austrobaileya 8(3): 225-251 (2011)

oblanceolate, narrowly obovate or elliptic, 8-
23 cm long, 2.5-7 cm wide, mostly acuminate,
sometimes acute or obtuse at apex, acute or
attenuate at base, glabrous or with scattered
hairs on midvein below; margins remotely
serrate towards tip or entire; secondary
veins 7—12(—15) pairs; petiole 3-10 mm long,
glabrous or pilose. Inflorescence a fascicle
of mostly 2-10 flowers, axillary or ramal;
bracts and bracteoles ovate to triangular,
sometimes denticulate, pubescent, persistent,
bracts 1.5-1.7 mm long, bracteoles 1-1.2 mm
long. Calyx limb 1.5-2 mm long; lobes ovate,
often denticulate and ciliate, 1-1.4 mm long,
pubescent. Petals oblong to obovate, 2.5-4
mm long, 1.4-1.8 mm wide, white or cream-
pink, glabrous or with a few hairs, sometimes
ciliolate. Stamens 15-30, 2.5-3.5 mm long,
filaments pilose. Disc pilose. Style 2.5-3 mm
long, shorter and without stigma in males,
sparsely pilose. Fruit cylindrical to narrowly
ellipsoidal, 15-25 mm long, 6-10 mm wide,
glabrescent, blue to purple.

Distribution and habitat : Endemic to the
Wet Tropics of Queensland where it occurs
from Big Tableland, north of Mt Finnigan,
south to the Mossman River, in mesophyll
and notophyll vine forest.

Two varieties are recognised.

Key to varieties

Young branchlets, petiole and lamina glabrous.var. cyanocarpa

Young branchlets, petiole and midvein of lamina pilose below.var. pilosa

7a. Symplocos cyanocarpa var. cyanocarpa

Shrub or small tree to 5 m. Shoots, young
branchlets, petiole and lamina glabrous
or nearly so. Bracts, bracteoles, and calyx
appressed puberulous. Fig. 2A-C.

Illustrations : Hyland et al. (2010).

Additional selected specimens examined : Queensland.

Cook District: Mt Finnigan, W slopes, Sep 1948, Brass
20043 (BRI); Cedar Bay N.P., W slopes of Mt Finnigan,
Oct 1999, Forster PIF25014 & Booth (BRI); Mt Misery,
Jun 1992, Forster PIF10751 et al. (BRI); Daintree N.P.,
Cedar Bay section. Gap Creek, Nov 2000, Forster
PIF26464 et al. (BRI); Cedar Bay N.P, Jul 1994, Forster
PIF15585 et al. (BRI); N of Daintree River, on road to

Cape Tribulation, Nov 1985, Tucker 37 (BRI); Palm
Road, 3 km W of Alexander Bay, Dec 2001, Forster
PIF27907 et al. (BRI); Whyanbeel, Dec 1978, Gray 1181
(BRI, L); loc. cit ., Dec 1978, Gray 1187 (BRI); c. 2 km
along Whyanbeel - Daintree track, from Whyanbeel
Creek, Jan 1986, Guymer 2060 (BRI); Near Cooper
Creek, Aug 1987, Sankowsky 639 & Sankowsky (BRI);
Mossman Gorge, W of Mossman, Jun 1972, Wrigley &
TelfordNQ1137 (BRI).

Distribution and habitat : Endemic to the
Wet Tropics of Queensland where it occurs
from the vicinity of Mt Finnigan to south of
Mossman (Map 6) in mesophyll and notophyll
vineforest on granitic and metamorphic
derived soils.

Jessup, Symplocos in Australia 237

Fig. 2. Symplocos cyanocarpa var. cyanocarpa. A. portion of stem with inflorescences and leaf base x 3. B. flower x
8. C. fruit x 2. A-C from Forster PIF10751 et al. (BRI). S. cyanocarpa var. pilosa. D. inflorescence and leaf base x
4, Hyland 10582 (BRI). S. glabra. E. branchlet with fruit x 0.8, Hyland 11798 (BRI). S. hayesii. F. inflorescence with
flowers x 4, Hyland 14628 (CNS). G. fruit x 2, Smith 10839 (BRI). H. leaf abaxial surface x 1, Smith 10839 (BRI). Del.
W. Smith.

238

Map 6. Distribution of Symplocos cyanocarpa var.
cyanocarpa

Phenology : Flowers are recorded from July
to January; fruit from November to March.

Conservation status : Symplocos cyanocarpa
var. cyanocarpa occurs mostly in National
Parks and other reserves throughout its range
and is not considered to be threatened.

7b. Symplocos cyanocarpa var. pilosa Jessup
varietas nova, differt a var. cyanocarpa
ramulis juvenibus, petiolo et costa abaxialiter
pilosis. Typus: Queensland. Cook District:
Timber Reserve 146, Tableland Logging
Area, 11 September 1980, B.P.Hyland 10582
(holo: BRI; iso: CNS).

Symplocos sp. (Big Tableland B.P.Hyland
10582); (Jessup 1997, 2002, 2007, 2010).

Illustrations : Hyland et al. (2010), under
Symplocos sp. (Big Tableland B.P.Hyland
10582).

Shrub or small tree to 3 m. Shoots, young
branchlets, petiole and main veins of leaves
on underside pilose with scattered erect hairs,
glabrous above. Bracts, bracteoles and calyx
pilose with more or less erect hairs. Fig. 2D.

Additional specimens examined : Queensland. Cook
District: T.R. 146, Tableland L.A., Jul 1975, Hyland
8324 (BRI, CNS); Hartley Creek at end of Mt Hartley
track, Oct 1996, Ford 1774 (BRI); Along track from
Hartley Creek to Mt Hartley, Oct 1996, Ford 1782 (BRI);
Mt Hartley, western slopes, T.R. 165, Jul 1995, Forster
PIF17296 & Figg (BRI).

Austrobaileya 8(3): 225-251 (2011)

Distribution and habitat : Endemic to the
Wet Tropics of Queensland where it occurs on
Big Tableland and Mt Hartley, northwest of
Cedar Bay (Map 7) in notophyll vineforest on
granite substrates.

Map 7. Distribution of Symplocos cyanocarpa var.
pilosa

Phenology : Flowers are recorded in July,
September and October and very immature
fruit in October.

Etymology : The varietal epithet is formed
from the Latin word pilosus (covered with
short, weak, thin hairs) and refers to the
indumentum on the young branchlets, petiole
and adaxial surface of the midvein.

Conservation status : Symplocos cyanocarpa
var. pilosa is only known from the localities
cited above. Additional collections are
required to determine its area of occupancy
and population numbers.

8. Symplocos gittinsii Jessup nom. et
stat. nov.; S. cochinchinensis var. gittonsii
Noot., Leiden Bot. Ser. 1: 162 (1975). Type:
Queensland. North Kennedy District: Mt
Spec Range, May 1962, C.H.Gittins 494 (holo:
BRI; iso: L).

Illustrations : Nooteboom (1982: Fig. 11);
Cooper & Cooper (2004: 537); Hyland et al.
(2010), all under Symplocos cochinchinensis
var. gittonsii

Tree to 30 m. Shoots and young branchlets
glabrous; cataphylls glabrous and ciliate.
Leaves coriaceous; lamina obovate,

239

Jessup, Symplocos in Australia

sometimes elliptic, 8-18 cm long, 3.5-9.5
cm wide, acuminate, acute or obtuse at apex,
acute or attenuate at base, glabrous; margins
entire, revolute; secondary veins mostly 7-
11 pairs; petiole 10-35 mm long, glabrous.
Inflorescence axes branched, to 13 cm long,
spicate, sparsely appressed puberulous; bracts
and bracteoles broadly or depressed ovate,
glabrous with ciliate margin, persistent to
after anthesis, sometimes to fruiting, bracts
0.8-1.5 mm long, bracteoles 0.7-1 mm long.
Calyx limb 0.7-1.5 mm long, glabrous; lobes
depressed obovate, mostly glabrous with
ciliate margin. Petals broadly obovate, 4-6
mm long, 3.5-4.5 mm wide, white, glabrous.
Stamens 60-70, 2.5-6 mm long. Disc slightly
raised, 5-lobed, glabrous. Ovary 3-locular,
with c. 3 ovules per locule. Style 4 mm long,
glabrous. Fruit ellipsoidal or ovoid, 6-7 mm
long, 4-5 mm wide, blue to black.

Additional selected specimens examined : Queensland.
Cook District: T.R. 30, Mt Lewis area, Oct 1967,
Hyland 1048 (BRI); Ridge between McKinnon &
Wrights Creeks, southwest of Edmonton, Jun 1996,
Forster PIF19263 et al. (BRI); S.F.R. 310, Gadgarra,
Feb 1981, Bragg s.n. (BRI [AQ874875]); T.R. 1230,
Boonjie F.A., May 1977, Gray 533 (BRI, MEF); S.F.R.
310, Gadgarra, Windin F.A., May 1990, Hyland 14017
(BRI); S.F.R. 310, Windin F.A., Jan 1978, Gray 850
(BRI, MEF); Topaz to Boonjee Road, May 1987, Starkey
74 (BRI); On Topaz Road, Dec 1985, Sankowsky 459 &
Sankowsky (BRI); Topaz Road, Nov 1992, Jessup 905
(BRI); Palmerston Highway, 1 km south-east of Beatrice
River, Jan 1993, Bean 5401 (BRI); Wooroonooran N.P.,
Palmerston section, Dec 1999, Forster PIF25209 (BRI);
S.F. 756, Mt Father Clancy, May 2000, Forster PIF25726
& Booth (BRI); Portion 297, Parish of Dirran, Apr 1984,
Gray 3992 (BRI, CNS). North Kennedy District: 4.5
km south along Culpa Road, S.F. 605 Koombooloomba,
Culpa F.A., May 2002, Forster PIF28705 & Booth
(BRI); Mt Spec near Paluma village. May 1985,
Jackes 14 (BRI); Paluma, May 1980, Horton s.n. (BRI
[AQ343157]); Paluma, Sep 1990, Gumming 10650 (BRI);
Mt Spec N.P, Paluma, behind Rangers barracks. May
1989, Fell DF1851 (BRI). South Kennedy District:
S.F. 652 Cawley, near Mt Macartney, Jun 1995, Forster
PIF16714 & Tucker (BRI); Clarke Range, 13.7km from
Eungella township, Eungella N.P, Apr 1991, Forster
PIF8057 & McDonald (BRI); Roadside c. 15 km N of
Eungella township on Dalrymple Road, Nov 1989,
McDonald 4465 (BRI); Dalrymple Heights, Eungella
Range, Fredrickson’s Farm, May 1987, Champion 227
(BRI).

Distribution and habitat : Endemic to
Queensland where it occurs from Mt Lewis
south to Culpa L.A. in the Wet Tropics, then
disjunct to Paluma and the Clarke Range near
Eungella (Map 8). Grows in upland notophyll

or mesophyll vine forest on diverse substrates
(basalts, granites, metamorphics, rhyolites).

Map 8. Distribution of Symplocos gittinsii

Phenology: Flowers are recorded in April and
May; fruit from November to January.

Notes: This species was treated as Symplocos
cochinchinensis var. gittonsii Noot. (as
S. cochinchinensis subsp. thwaitesii var.
gittonsii ) in Nooteboom (1982).

Etymology: The name commemorates
Clifford Halliday Gittins (1904-1995),
an amateur botanical collector of mostly
Queensland plant specimens.

Conservation status: Symplocos gittinsii
occurs in several National Parks and other
reserves throughout its range and is not
considered to be threatened.

9. Symplocos glabra Jessup nom. et stat.
nov.; S. cochinchinensis var. glaberrima
Noot., Brunonia 4: 319-320 (1982). Type:
Queensland. Cook District: State Forest
Reserve 194, Compartment 54, 17 March
1971, G.C. Stocker 622 (holo: L; iso: BRI,
CNS, MEL).

Illustrations: Nooteboom (1982, Fig. 11);
Hyland et al. (2010), both under Symplocos
cochinchinensis var. glaberrima.

Small tree to 9 m. Shoots and young
branchlets glabrous. Leaves coriaceous;
lamina oblanceolate or narrowly obovate,
rarely elliptic, 7-15 cm long, 2-7 cm wide,
acuminate, acute or obtuse at apex, acute or
attenuate at base, glabrous; margins entire,

240

Austrobaileya 8(3): 225-251 (2011)

revolute; secondary veins mostly 8-12 pairs;
petiole 3-10 mm long, glabrous. Inflorescence
axes 2 or 3, racemose, to 3 cm long, glabrous,
branching near base, sometimes single;
pedicels 0.2-2 mm long; bracts and bracteoles
glabrous but ciliate, caducous or persistent to
anthesis, bracts broadly ovate or oblong, 1.2-
2.5 mm long, bracteoles ovate, 0.7-1.2 mm
long. Calyx limb 1.2-1.5 mm long, glabrous;
lobes depressed obovate, glabrous with ciliate
margin. Petals narrowly obovate, 4.5-6 mm
long, 2-3 mm wide, cream to white, glabrous.
Stamens mostly 30-50, 2-5 mm long. Disc 5-
lobed, sparsely puberulous. Style 4.5-5 mm
long, glabrous. Fruit subglobose or ellipsoidal,
8-11 mm long, (5-) 8-10 mm wide, blue-black
to purple-black. Fig. 2E.

Additional selected specimens examined : Queensland.
Cook District: V.C.L. Noah, Mt Hemmant, Oct 1975,
Hyland RFK3339 (BRI); Mt Windsor F.R, c. 5.7 km
from Spencer Creek Forestry barracks, Jun 2004, Ford
4341 & Hewett (BRI); S.F.R. 144 Whypalla, Bower Bird
L.A., Dec 1987, Hyland 13454 (BRI); Mt Spurgeon, W of
Mossman, Sep 1936, White 10557 (BRI); Mt Bellenden
Ker, Jun 1969, Smith 14682B (BRI); Wooroonooran
N.R, Mt Bellenden Ker summit area, Dec 2000, Forster
PIF26510 et al. (BRI); loc. cit ., Dec 2001, Forster
PIF27945 et al. (BRI); Mt Bellenden Ker, c. 0.5 miles
[0.8 km] SW of centre peak, Jun 1969, Smith 14638 &
14638a (BRI); Summit of Centre Peak, Bellenden Ker,
Nov 1972, Webb & Tracey 11920 (BRI); S.F.R. 194
Herberton, c. 10 miles [16 km] S of Atherton, May 1964,
Rudder s.n. (BRI [AQ84400], L); Bellenden Ker, Jan
1923, White s.n. (BRI [AQ84393]); S.F.R. 310 Bellenden
Ker Range, Jan 1980, Hyland 10205 (BRI); Mt Bellenden
Ker, Feb 1982, Irvine 2210 (BRI, CNS); Mt Bellenden
Ker, Dec 2000, Jago 5779 & Forster (BRI); Summit of
Mt Bellenden Ker, Aug 1971, Hyland5312 (BRI); S.F.R.
755 Gosschalk L.A., Jun 1982, Hyland 11798 (BRI); 100
m W of summit of NW peak of Bartle Frere, Jul 1995,
Hunter JH4461 (BRI); Wooroonooran N.P., summit of
South Peak, Oct 1997, Forster PIF21771 et al. (BRI);
Mt Bartle Frere, Jun 1961, Martin & Hyland AFO/1890
(BRI); S.F.R. 194 Barron, Jan 1983, Gray 2909 (BRI);
S.F.R. 251, Ravenshoe - Tully Falls Road, Aug 1968,
Hyland 1737 (BRI).

Distribution and habitat: Endemic to the Wet
Tropics of Queensland where it occurs at Mt
Hemmant, Mt Windsor Tableland, Mt Carbine
Tableland, on the Lamb Range, Bellenden Ker,
Cardwell and Herberton Ranges (Map 9) in
simple to complex microphyll and notophyll
vineforests on granites (upper montane
localities) and rhyolites.

Phenology: Flowers are recorded from
December to May; fruit from December to
March.

Note: The superlative epithet ‘glaberrima’ is
not available for use at the rank of species.
This species was treated as Symplocos
cochinchinensis var. glaberrima Noot. (as
S. cochinchinensis subsp. thwaitesii var.
glaberrima) in Nooteboom (1982).

Conservation status: Symplocos glabra
occurs in several National Parks and other
reserves in the Wet Tropics and is not
considered to be threatened.

Etymology: The specific epithet is derived
from the Latin adjective glaber (glabrous) and
refers to the lack of indumentum on all parts
of the plant.

10. Symplocos graniticola Jessup,
Austrobaileya 4: 9 (1993) Type: Queensland.
Cook District: Mt Lewis, 24 December 1986,
G. Sankowsky 598 & N. Sankowsky (holo:
BRI).

Symplocos sp. ‘Mt Lewis’; (Thomas &
McDonald 1989).

Illustrations: Jessup (1993: 10); Hyland et al.
( 2010 ).

Shrub to 3 m. Shoots and cataphylls with
very sparse appressed hairs, glabrescent;
cataphylls caducous. Twigs glabrous. Leaves
coriaceous; lamina narrowly obovate or
narrowly oblanceolate, obtuse or shortly
acuminate, 9-23 cm long, 2-7.5 cm wide,
glabrous; margins entire, recurved, sometimes
remotely serrulate or denticulate, acute or

241

Jessup, Symplocos in Australia

attenuate at base; secondary veins mostly
6-12 pairs; petiole 8-20 mm long, glabrous.
Inflorescence axes clustered, spicate, to 1.5
cm, puberulous; bracts and bracteoles ovate,
slightly keeled, sometimes denticulate,
pubescent, persistent, bracts 1.2-1.5 mm long,
bracteoles 1—1.3 mm long. Calyx limb tube 1-
1.3 mm long, glabrous; limb 1-1.2 mm long;
lobes ovate or depressed ovate, 0.8-1 mm
long, sparsely puberulous to nearly glabrous
and with a ciliate margin. Petals oblong to
obovate, 2.5-3.5 mm long, 1.5-2 mm wide,
white, glabrous. Stamens 25-30, 2.5-4 mm
long, filaments glabrous. Disc pilose. Style
2.5 mm long, pilose. Fruit ovoid or ellipsoid,
10-18 mm long, 6-12 mm wide, black.

Additional selected specimens examined : Queensland.

Cook District: Platypus Creek, Jan 1936, Flecker 1216a
(CNS); Pinnacle Rock track, Jun 1992, Forster PIF10716
etal. (BRI); North Mary L.A., S.F. 143,31.5 km along Mt
Lewis road, Jun 1995, Forster PIF16774A (BRI); S.F.R.
143, South Mary L.A., May 1976, Hyland 8773 (BRI,
CNS); 32.6 km along Mt Lewis Road from Mossman
- Mt Molloy Road, Dec 1989, Jessup GJD3359 et al.
(BRI). S.F.R. 143, Kanawarra, Carbine Tableland, Nov
1995, Gray 6400 (BRI); Mt Lewis F.R., off Mt Lewis
Road, Jan 2008, FordAF5188 et al. (BRI).

Distribution and habitat : This species is
endemic to the Wet Tropics of Queensland
where it is known from Mt Spurgeon to Mt
Lewis, Platypus Creek and Pinnacle Rock
near Mossman (Map 10). It grows in simple
microphyll and notophyll vineforests on
granite or granodiorite substrates.

Map 10. Distribution of Symplocos graniticola

Phenology : Flowers are recorded from
December to May; green fruit from June to
September, mature fruit in November.

Conservation status: Symplocos graniticola
is currently listed as Vulnerable under the
NCA.

11. Symplocos harroldii Jessup,
Austrobaileya 4: 7 (1993). Type: Queensland.
Moreton District: Moggill State Forest, just
downstream of road crossing Pullen Creek, 16
November 1980, L.W.Jessup 268 (holo: BRI;
iso: K, NSW).

Symplocos sp. 1.; Stanley & Ross (1986).

Illustrations: Jessup (1993: 8); Leiper et al.
(2008: 336).

Shrub or small tree to 6 m. Shoots and twigs
pilose. Leaves coriaceous; lamina narrowly
obovate or oblanceolate, 3-10 cm long, 0.8-
3.5 cm wide, acuminate at apex, acute at base,
glabrous or with a few hairs on underside,
particularly on midvein; margins denticulate
to serrate towards tip; secondary veins mostly
6-12 pairs; petiole 3-4 mm long, usually
with scattered hairs. Inflorescence racemose,
to 2 cm, puberulous; pedicels 0.2-2 mm
long; bracts and bracteoles ciliate, semi-
persistent, bracts broadly triangular, 0.7-1
mm long, bracteoles triangular, 0.3-0.4 mm
long. Hypanthium glabrous. Calyx lobes
broadly ovate, 0.5-0.7 mm long, glabrous,
often with a ciliate margin. Petals obovate to
oblong, 3-4 mm long, 2-2.5 mm wide, white,
glabrous. Stamens 30-40, 1.8-5 mm long.
Disc puberulous. Ovary 3-locular with up
to 4 ovules per locule. Style 3.5-4 mm long,
glabrous, dilated at stigma. Fruit narrowly
ovoid or ellipsoid, 8-10 mm long, 6-7 mm
wide, red.

Additional selected specimens examined: Queensland.
Wide Bay District: Tinana Creek, 7 km E Tiaro, Dec
1992, Forster PIF12503 & Smyrell (BRI); Tinana
Creek, 7 km ENE of Tiaro, Mar 1992, Smyrell s.n. (BRI
[AQ541786]); S.F. 502, Veteran L.A., Gympie, Sep 1992,
Thomas 8 & Barry (BRI); 1 km ENE of Timberwah
[Tinbeerwah] Hall, junction forestry road & tributary of
Wooroi Creek, S.F. 959, Sep 1992, Thomas 1 & Barry
(BRI); Gum Tree Drive off Cooroy - Tewantin Road,
Sep 1992, Thomas 5 & Barry (BRI); Noosa N.P., off
Viewland Drive, Jan 1993, Thomas & Gayle s.n. (BRI
[AQ636338]); Panorama Drive via Duke Road, 8 km SE
of Cooroy, Apr 1993, Bean 5928 (BRI); N of Ringtail

242

Creek Road, Noosa Shire, Portion 8V, near S.F. 997,
Oct 1994, Thomas 102 (BRI); Noosa Shire hinterland
N of Collwood Road, near Lake McDonald, Lot 3 Rp
800331 Parish ofTewantin, Sep 1995, Thomas 33 (BRI);
Compartment 1, Ringtail S.F., NE of Pomona, Sep
1997, Bean 12412 (BRI); Doonan Wetland Conservation
Area, Jan 2007, Thomas MS112 (BRI); Maroochy Shire
Bushland Conservation Reserve, Doonan Creek East
(BCR), Jul 2007, Thomas MS139 (BRI); Near Doonan
Creek, Feb 2002, Thomas 420M (BRI). Moreton
District: 650 Campbells Pocket Road, Caboolture, Jan
2006, Weber s.n. (BRI [AQ735727]); 0.5 km SW of
McAfees Lookout, S.F. 309, Feb 1991, Forster PIF7790
& Bird (BRI); Dillon Road, The Gap c. 0.5 km from
intersection with Payne Road, Jul 1999, Harris WKH124
& Thompson (BRI); Greenford Street, Chapel Hill, Sep
1999, Stephens s.n. (BRI [AQ664972]); Cubberla Creek,
Fig Tree Pocket, Sep 1998, Stephens 2 (BRI); Daisy Hill
S.F., Logan City, Dec 1993, Thompson MOR323 (BRI);
Hinze Dam, Mar 2007, Jinks s.n. (BRI [AQ784176]);
Hinze Dam, c. 3.5 km SW of Mudgeeraba, Apr 2007,
Jinks DBA { BRI).

Distribution and habitat : This species is
endemic to southeast Queensland. Collections
of this species made since it was described in
1993 have extended the known range from
Tinana Creek near Tiaro to the Hinze Dam
near Mudgeeraba (Map 11).

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Map 11. Distribution of Symplocos haroldii

Austrobaileya 8(3): 225-251 (2011)

Phenology : Flowers are recorded from
September to November; fruit in January.

Note : The phrase in the description in Jessup
(1993) stated ‘peduncles 0.2-2 mm long’; this
should have read ‘pedicels 0.2-2 mm long’.

Conservation status : Symplocos harroldii is
currently listed as Near Threatened under the
NCA.

12. Symplocos hayesii C.T.White & Francis,
Proc. Roy. Soc. Queensland 33: 158, t.2 (1921).
Type: Queensland. Cook District: Glenallan
[Glen Allyn], Malanda, s.dat., H.C.Hayes
(holo: BRI [AQ23234]).

Illustrations: Cooper & Cooper (2004: 538);
Hyland et al. (2010).

Shrub or small tree to 2 m. Shoots and young
branchlets with persistent cataphylls, pilose.
Leaves chartaceous; lamina narrowly ovate
or narrowly obovate or elliptic, 6-12 cm long,

2.5- 5.5 cm wide, acuminate at tip, rounded
or cordate at base, glabrous except midvein
on underside; margins serrate; secondary
veins mostly 7-10 pairs; petiole 3-7 mm long,
pilose. Inflorescence spicate, to 1.5 cm, or a
fascicle, pilose; bracts and bracteoles sparsely
pilose and ciliate, persistent, bracts ovate to
narrowly triangular, 3-9 mm long, bracteoles
ovate, 2-3 mm long. Calyx limb 1-1.5 mm
long, glabrous; lobes ovate, acute, glabrous or
with a few hairs, margin often ciliate. Petals
ovate or oblong, denticulate, 2.2-2.5 mm
long, white, glabrous. Stamens mostly 12-15,
as long as petals in males, shorter in females.
Disc pilose. Style 1.5-2 mm long in females,
glabrous, stigma peltate. Fruit narrowly ovoid
to ellipsoidal, usually curved, 15-20 mm long,

3.5- 5 mm wide; blue to black, mesocarp thin,
fleshy; endocarp smooth; seed 1. Fig. 2F-H.

Additional selected specimens examined : Queensland.
Cook District: S.F. 185 Danbulla, 1 km N of Hoop Pine
Triangle on C road, Jul 1994, Forster PIF15609 et al.
(BRI); Danbulla, 12 miles [19.3 km] NE of Atherton, Jun
1949, Smith 4181 (BRI); Yungaburra, near Lake Barrine,
Blake 9602 (BRI, CANB); near Lamond’s Hill, 7.5 miles
[12 km] SE of Malanda, Sep 1959, Smith 10839 (BRI);
S.F.R. 310, Gadgarra, Boonjee L.A., Jan 1993, Hyland
14628 (CNS); T.R. 1230, Boonjee L.A., Gray 1246 (BRI);
S.F. 756, Mt Father Clancy, May 2000, Forster PIF25720
& Booth (BRI); Vacant Crown Land, Bartle Frere, E of
Glen Allyn Trig., Webb & Tracey 5795 (BRI). North
Kennedy District: tributary of Chunum Creek, May
2005, Wannan 3986 & Sinclair (BRI); Tableland L.A.,

243

Jessup, Symplocos in Australia

0.5 km S of Koolmoon Creek near junction of Ebony
Road and Tully Falls Road, 6 km NW of Tully Falls, Oct
1998, Jessup GJM2670 et al. (BRI).

Distribution and habitat : Endemic to the
Wet Tropics of Queensland where it occurs
from Tinaroo Range to Koolmoon Creek,
southeast of Ravenshoe (Map 12). Grows
in mesophyll and notophyll vineforests on
substrates derived from basalts, granites and
metamorphics.

Phenology: Flowers are recorded from
December to February; fruit from May to
September.

Conservation status: Symplocos hayesii
occurs in several National Parks and other
reserves on the Atherton Tableland is not
considered to be threatened.

13. Symplocos hylandii Noot., Brunonia 4:
323 (1982). Type: Queensland. Cook District:
State Forest Reserve 755 Boonjee, 21 March
1968, B.HylandRFK1395 (holo: CNS).

Illustrations: Hyland et al. (2010).

Shrub or small tree to 3 m. Shoots and
young branchlets with persistent cataphylls,
pilose. Feaves coriaceous; lamina narrowly

obovate, 10-16 cm long, 1.7-3.5 cm wide,
often bullate and brittle, acuminate at apex,
acute at base, glabrous or with a few hairs
on underside, particularly midvein; margins
serrate; secondary veins mostly 5-12 pairs,
raised below, sunken above; petiole 3-5 mm
long, pilose. Inflorescence spicate, to 1.5 cm,
pilose; bracts and bracteoles ovate, sparsely
pilose and ciliate, persistent, bracts, c. 2 mm
long, bracteoles, c. 1 mm long. Calyx limb
1.3-1.7 mm long, glabrous; lobes ovate or
broadly oblong, puberulous or nearly glabrous,
margins entire or minutely denticulate. Petals
ovate or oblong, c. 3 mm long, white, glabrous.
Stamens 10-15. Disc pulvinate, glabrous. Fruit
ampulliform to ovoid, (7-) 10-12 mm long,
8-9 mm wide, blue, mesocarp thin, endocarp
shallowly grooved; seed 1. Fig. 3A,B-

Additional selected specimens examined : Queensland.
Cook District: Mt Bellenden Ker, c. 7/8 mile [1.5 km]
SE of centre peak, Jun 1969, Smith 14717 (BRI); East
Mulgrave River Falls, May 2002, Jago 6212 (BRI);
T.R. 1230, Stockwellia Track, 4.9 km from Lamins Hill
turnoff (headwaters of Russell River), Aug 1989, Bostock
912 & Guymer (BRI); Swipers Flat, WooroonooranN.R,
Jul 1995, ForsterP1F17136& F/gg/BRI); Wooroonooran
N.R, track to Chuck Lunga Creek, E of Boonjee, Jun 1996,
Forster PIF19284 & Tucker (BRI); Wooroonooran N.R,
lower reaches of Russell River Track, Jan 2007, Ford
AF4913 & Metcalfe (BRI); N.P.R. 904 Wooroonooran,
Coolamon Creek, Oct 2001, Ford AF3047 et al. (BRI);
Coolamon Creek, end of Towalla Road, Wooroonooran
N.R, Oct 2001, Cooper WWC1606 (BRI).

Distribution and habitat: This species is
endemic to the Wet Tropics of Queensland
where it occurs from Mt Bellenden Ker
to near Millaa Millaa (Map 13) mostly in
mesophyll vineforests on substrates derived
from granites and metamorphics.

Phenology: Flowers are recorded in May;
fruit from October to January.

Conservation status: Symplocos hylandii is
located mostly in Wooroonooran N.P and is
not considered to be threatened.

244

14. Symplocos oresbia Jessup species nova

S. wooroonooran simulans sed innovationibus
et axibus inflorescentiarum glabris, bracteis
et bracteolis ante anthesin caducis, petalis
2-2.5 mm longis (non 4-5 mm) et 1-1.3 mm
latis (non 1.4-2.5), staminibus paucioribus
(16-19 non 30-35), stylis brevioribus (1.5-1.8
mm non 3-3.5 mm) et fructibus amplissimus
(10-15 x 7-9 mm non 5.8-6.3 x 4-5.5 mm)
differt. Typus: Queensland. Cook District:
Summit area of Thornton Peak, 12 November
1973, TG.Hartley 14045 (holo: BRI; iso:
CANB).

Symplocos sp. (Mt Finnigan L. J.Brass 20129);
(Jessup 1994, 1997, 2002, 2007, 2010).

Illustrations : Hyland et al. (2010), under
Symplocos sp. (Mt Finnigan L. J.Brass
20129).

Shrub or small tree to 8 m. Shoots and young
branchlets glabrous. Leaves coriaceous;
lamina elliptic, 3-8 cm long, 1-3 cm wide,
acuminate at apex, acute or attenuate at base,
glabrous; margins entire, revolute; secondary
veins mostly 7-9 pairs; petiole 3-7(-12) mm
long, glabrous. Inflorescence axes 2 or 3,
racemose, to 3 cm long, glabrous, branching

Austrobaileya 8(3): 225-251 (2011)

near base, sometimes single; pedicels 0-0.2
mm long, bracts broadly ovate, 1-1.2 mm long,
bracteoles ovate, 0.5-0.6 mm long, glabrous
or ciliate, caducous before anthesis; pedicel
0.2-1.5 mm long. Calyx limb 1-1.5 mm long,
lobes depressed ovate or rounded, 0.5-0.7
mm long, glabrous with ciliate margin. Petals
obovate, 2-2.5 mm long, 1-1.3 mm wide,
rounded, ?white, glabrous. Stamens 16-19;
filaments glabrous. Disc glabrous. Style
1.5-1.8 mm long, glabrous; stigma indistinct.
Fruit ampulliform, 7.8-8.5 mm long, 4.6-5.5
mm wide, colour unknown. Fig. 3C,D.

Additional selected specimens examined : Queensland.
Cook District: Mt Finnigan, Apr 1948, Brass 20129
(BRI, CANB); Cedar Bay N.P., Mt Finnigan summit
area, campsite on Horans Creek, Oct 1999, Forster
PIF25035 & Booth (BRI); Summit of Devils Thumb,
Mossman Gorge N.P., Apr 1986, Godwin C3093 (BRI);
Pinnacle Rock Track, Whyanbeel, Feb 1996, Gray 6566
(BRI); Mossman Bluff, about 10 km W of Mossman,
Dec 1988, Fell DF1686 (BRI).

Distribution and habitat : This species is
endemic to the Wet Tropics of Queensland
where it occurs from Mt Finnigan to Mossman
Bluff (Map 14) in exposed montane rainforest
(microphyll to nanophyll vineforests and
vinethickets) on substrates derived from
granite.

Map 14. Distribution of Symplocos oresbia

Phenology : Flowers are recorded from
February to April; fruit from December to
January.

Notes: Specimens of this species (e.g. Brass
20129 ) were included under Symplocos
cochinchinensis subsp. thwaitesii var.
montana in Nooteboom (1982).

Conservation status: Symplocos oresbia (as
Symplocos sp. (Mt Finnigan L. J.Brass 20129))
is currently listed as Near Threatened under
the NCA.

Jessup, Symplocos in Australia

245

Fig. 3. Symplocos hylandii. A. node with leaf base and budding inflorescence x 4, Jago 6212 (BRI). B. fruit x 2, Ford
AF4913 & Metcalfe (BRI). S. oresbia. C. flower bud x 8, Godwin C3093 (BRI). D. leaf base and inflorescence with
fruit x 2, Hartley 14045 (BRI). S. wooroonooran. E. node with leaf base and inflorescence with buds and flowers x
3, Forster PIF21767 et al. (BRI). F. Flower x 6, Forster PIF21767 et al. (BRI). G. fruit x 3, Hyland 5775 (BRI). Del.
W. Smith.

15. Symplocos paucistaminea F.Muell. &
F.M.Bailey ex F.M.Bailey, Syn. Queensland
FI, Supp. 3: 46 (1890). Type: Queensland.
Cook District: Harvey’s Creek, in 1889
(Bellenden Ker Expedition), F.M. Bailey
s.n. (holo: BRI [AQ23235]; iso: BM, K, L,
MEL46497).

Illustrations : Cooper & Cooper (2004: 538);
Hyland et al. (2010).

Dioecious shrub or tree to 18 m. Shoots and
young branchlets reddish-brown to dark brown
pilose, cataphylls rarely evident. Leaves thinly
coriaceous or chartaceous; lamina narrowly

246

Austrobaileya 8(3): 225-251 (2011)

obovate, oblanceolate or elliptic, 5-22 cm
long, 2-8.5 cm wide, acuminate at apex,
acutely or obtusely cuneate or rounded at
base, glabrescent above, pilose below, slightly
glossy; margins serrulate and denticulate;
secondary veins mostly 7-12 pairs, depressed,
flush or indistinct on upper surface; petiole 4-
12 mm long, pilose. Inflorescences clustered
or not, spicate, to 5 cm, pilose; bracts and
bracteoles ovate, reddish-brown pilose or
villous, persistent, bracts 1.5-2.5 mm long,
bracteoles 1.2-1.5 mm long. Calyx limb 1.2-

1.5 mm long; lobes narrowly ovate or oblong,
1-1.2 mm long, glabrous or sparsely pubescent
towards apex. Petals obovate, 2.5-3.5 mm
long, 1.3-1.5 mm wide, cream to white,
mostly glabrous. Stamens 15-20 in males,
7-10 in females and apparently sterile, 1-2.5
mm long. Disc glabrous. Style 1.2-1.8 mm
long, glabrous; females with peltate stigma.
Fruit ampulliform (when dry), globular when
fresh, 5-5.8 mm long, 3.5-4 mm wide, dark
blue to purple.

Additional selected specimens examined : Queensland.

Cook District: Mt Cook, Aug 1985, Scarth-Johnson
1641A (BRI); Gap Creek, T.R. 146, Fritz L.A., Jul 1973,
Stocker 997 (BRI); Cooper Creek, Cape Tribulation
Area, Aug 1996, Telford 12207 (BRI); V.C.L. Noah, Jun
1975, Hyland 8291 (BRI); Portion 188, Alexandra, Palm
Road, Jun 1985, Gray 4074 (BRI); Baileys Creek area, c.
0.2 miles [0.3 km] E of sawmill, c .7.5 miles [12 km] ENE
of Daintree, Oct 1962, Smith 11516 (BRI); Mt Spurgeon,
Apr 1968, Hyland RFK1508 (BRI); Nissen Creek, Mt
Perseverance Road, Jul 1995, Forster PIF17169 & Figg
(BRI); Churchill Creek, Churchill L.A., S.F. 143, Jul
1995, Forster PIF17204 & Figg (BRI); Churchill Creek,
S.F. 143, Churchill L.A., Nov 1998, Forster PIF24008 et
al. (BRI); F.R. 185, Kalorama L.A. near Danbulla, Jun
1961, Hyland 2047 (BRI); T.R. 1230, Boonjie L.A., May
1977, Gray 526 (BRI); Wooroonooran N.P., Palmerston
section. South Johnston Forestry Track., May 2005,
Forster PIF30970 & Jensen (BRI); S.F. 756 Mt Father
Clancy, May, 2000, Forster PIF25723 & Booth (BRI);

5.6 km S of Mena Creek township, SSW of Innisfail, Jul
2007, Bean 26546 (BRI). North Kennedy District: 27.5
km south along Culpa Road, S.F. 605 Koombooloomba,
May 2002, Forster PIF28789 & Booth (BRI); Paluma
Range, Dotswood Holding, Jun 1974, Hyland 3001
(BRI). South Kennedy District: Mt Macartney, S.F. 652
Cawley, May 1992, Forster PIF9911 & Tucker (BRI);
Dalrymple Heights and vicinity, Jun-Nov 1947, Clemens
s.n. (BM, BRI [AQ84447]).

Distribution and habitat: This species occurs
from the northern end of the Wet Tropics at
Cooktown south to the Clarke Range near
Eungella (Map 15). It grows in upland and

Map 15. Distribution of Symplocospaucistaminea

lowland mesophyll and notophyll vineforests
on substrates derived from basalts and
metamorphics.

Phenology: Flowers are recorded from May to
August; fruit from September to December.

Conservation status: Symplocos

paucistaminea is widely distributed in north
and central eastern Queensland and is not
considered to be threatened.

16. Symplocos puberula Jessup nom. et stat.
nov.; S. cochinchinensis var. pilosiuscula
Noot., Brunonia 4: 320 (1982). Type:
Queensland. Cook District: Speewah,
Upper Clohesy River, 5 June 1966, W.Rijkers
NQNC14754 (holo: L; iso: BRI, CNS).

Illustrations: Nooteboom (1981: Fig. 11);
Hyland et al. (2010), both under Symplocos
cochinchinensis var. pilosiuscula.

Tree to 25 m. Shoots and young branchlets
appressed puberulous, young branchlets
glabrescent. Leaves coriaceous; lamina
oblanceolate or obovate, sometimes elliptic,
6-15 cm long, 2.5-6 cm wide, acuminate
at apex, acute at base, glabrous or with a
few hairs on midvein on underside; margins
shallowly recurved, denticulate, nearly entire;
secondary veins mostly 6-12 pairs; petiole 5-
17 mm long, sparsely appressed puberulous,

247

Jessup, Symplocos in Australia

glabrescent. Inflorescence paniculate or
racemose to 7 cm, puberulous; pedicels 0.6-2
mm long; bracts and bracteoles pubescent,
caducous, bracts ovate, c. 1.5 mm long,
bracteoles narrowly triangular, c. 0.8 mm long.
Calyx limb 1-1.5 mm long, glabrous; lobes
depressed ovate or rounded, glabrous with
ciliate or denticulate margin. Petals oblong
or narrowly obovate, 3.5-5 mm long, 1.7-3.5
mm wide, white, glabrous. Stamens mostly
50-70, 2-6 mm long. Disc annular, shallowly
lobed, glabrous or with a few hairs. Style 3-4
mm long, glabrous or with a few hairs at base,
dilated at stigma. Fruit ampulliform, 7-10
mm long, 4-6 mm wide, dark blue to black.

Additional selected specimens examined : Queensland.
Cook District: 33 km along Leo Creek road, Mcllwraith
Range, Jun 1995, Forster PIF16822 (BRI); T.R. 14,
Head of Peach Creek, Mcllwraith Range, 26.5 km NE
of Coen, Cape York Peninsula, Aug 1993, Fell DGF3543
& Jensen (BRI); Hopevale Aboriginal Reserve, 13.2
km ESE of Hopevale Community, Endeavour River
right arm, Nov 1993, Fell DGF3859 & Stanton (BRI);
Mt Poverty Mine Road, near Normanby Tin Mines, Jun
2003, McDonald KRM1447 (BRI); Gap Creek, Cedar
Bay N.P., Jun 1992, Forster PIF10732 et al. (BRI);
Daintree N.P., Adeline Creek headwaters, top of hill
929, May 1999, Forster PIF24538 & Booth (BRI); S.F.
144, Mt Windsor Tableland, May 1979, Moriarty 2614
(BRI); T.R. 66, 3 km along Mt Lewis Road, Jun 2000,
Forster PIF25821 & Booth (BRI); S.F.R. 607, Shoteel
L.A., Oct 1982, Gray 2779 (BRI); S.F.R. 933, Little Pine
L.A., Feb 1983, Gray 2963 (BRI); Longlands Gap, S.F.

Map 16. Distribution of Symplocos puberula

194, Sep 2001, Forster PIF27514 et al. (BRI). North
Kennedy District: S.F.R. 605, Hilton L.A., Sep 1979,
Dockrill 1615 (BRI); Goold Island [Goold Island N.P],
Aug 1997, Cumming 16252 (BRI); Conway S.F., 2 km E
of Impulse Creek, along Timber Road, Sep 1994, Perry 2
(BRI); S.F. 299 Conway, Brandy Creek Road, 8 km S of
Airlie Beach, Feb 2004, Forster PIF29974 et al. (BRI).
South Kennedy District: 1 km SW of Mt Macartney,
Kangaroo Creek, Cathu S.F., Aug 1989, Thompson 148
(BRI); St Helens Creek, Eungella N.P., Aug 1992, Bean
4949 (BRI); Sarina Range, East Funnel Creek Road,
Sarina Shire, May 2006, Von Fahland 2 (BRI).

Distribution and habitat: This species occurs
from the Mcllwraith Range on Cape York
Peninsula south to the Sarina Range south
of Sarina (Map 16). Grows in mesophyll and
notophyll vineforests on substrates derived
from granites and metamorphics.

Phenology: Flowers are recorded from May to
November; fruit from December to February.

Notes: This species was treated as Symplocos
cochinchinensis var. pilosiuscula Noot. (as
S. cochinchinensis subsp. thwaitesii var.
pilosiuscula Noot.) in Nooteboom (1982).

Conservation status: Symplocos puberula is
widely distributed from Cape York Peninsula
to central eastern Queensland and is not
considered to be threatened.

17. Symplocos stawellii F.Muell., Fragm.
5: 60 (1865); S. stawellii var. stawellii ,
C.T.White, Proc. Roy. Soc. Queensland 50:
82 (1939); S. cochinchinensis var. stawellii
(F.Muell.) Noot., Leiden Bot. Ser. 1: 161
(1975). Type: Queensland. Rockingham Bay,
s.dat., J.Dallachy s.n. (syn: MEL).

Symplocos spicata var. australis Benth., FI.
Austral. 4: 292 (1868). Type: Queensland.
Rockingham Bay, s.dat. J.Dallachy s.n. (syn:
MEL); New South Wales. Richmond River,
s.dat., C. Moore s.n. (syn: NSW).

Illustrations: Nooteboom (1982, Fig. 11);
Hyland et al. (2010), both under Symplocos
cochinchinensis var. stawellii ; Floyd (1989:
397); Leiper et al. (2008: 336) both as S.
stawellii.

Shrub or tree to 30 m. Shoots sparsely
appressed pubescent, glabrescent; young
branchlets glabrescent. Leaves coriaceous;
lamina elliptic to oblanceolate or obovate,
6-18 cm long, 2-6 cm wide, acuminate
at apex, mostly acute or attenuate at base,

248

glabrous or with a few hairs on midvein on
underside; margins serrulate and crenulate,
sometimes slightly recurved; secondary veins
mostly 7-10 pairs; petiole 6-25 mm long,
glabrescent or glabrous. Inflorescence axes
branched, spicate, to 6 cm long, pubescent;
bracts and bracteoles broadly or depressed
ovate, sparsely pubescent or glabrous, ciliate,
persistent to fruiting, bracts 0.8-1.5 mm long,
bracteoles 0.6-0.8 mm long. Calyx limb 0.3-
0.5 mm long, glabrous; lobes very shallowly
rounded or depressed ovate, 0.1-0.4 mm
long, glabrous or nearly so and with a ciliate
margin. Petals obovate, 3.2-3.5 mm long,
1.7-2.5 mm wide, white, glabrous. Stamens
mostly 30-50, 1.5-3 mm long. Disc 5-lobed,
glabrous. Style 2.5-3.5 mm long, glabrous.
Fruit mostly ovoid, 5-5.5 mm long, 3.5-4.5
mm wide, blue to black.

Additional selected specimens examined : Queensland.
Cook District: Lake Barrine, Atherton Tableland, Sep
1929, Kajewski 1237 (BRI, L, P); Malanda - Atherton
Road, May 1976, Stocker 1438 (BRI, CNS, L). North
Kennedy District: S.F.R. 251, Parish of Ravenshoe, Jun
1988, Gray 4844 (BRI, CNS). South Kennedy District:
Dalrymple Road, Eungella N.P, Apr 2000, Forster

7-

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u

T-:

4 Rockham

V

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pton

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Map 17. Distribution of Symplocos stawellii

Austrobaileya 8(3): 225-251 (2011)

PIF25512 & Booth (BRI); S.F.R. 679, Range L.A., 1 km
S of Crediton, Clarke Range, Jun 2001, Ford AF2887
(BRI). Port Curtis District: Byfield N.P., 9 km ENE
of Byfield township, Sep 2000, Forster PIF26237 et al.
(BRI). Wide Bay District: Verrierdale Road, 8 km W of
Peregian Beach, May 1990, Bean 1589 (BRI). Moreton
District: Running Creek, c. 2 miles [3.2 km] NW of
Yandina, May 1969, Smith 14616 (BRI). New South
Wales. Coombadjha Creek Road, Apr 1980, Floyd 1477
(BRI); Johns River near Newcastle, May 1956, Webb &
Tracey 3691 (BRI).

Distribution and habitat : This species is
widespread in eastern Queensland and New
South Wales from the Hann Tableland,
northwest of Mareeba to near Windsor, New
South Wales (Map 17). It generally grows in
notophyll vineforests, although it sometimes
occurs as an understorey tree in moist eucalypt
forest. Substrates are diverse.

Phenology : Flowers are recorded from April
to August; fruit from November to January.

Conservation status : Symplocos stawellii is
widely distributed from north Queensland to
central eastern New South Wales and is not
considered to be threatened.

18. Symplocos thwaitesii F.Muell., Fragm.
3: 22 (1862); S. cochinchinensis subsp.
thwaitesii (F.Muell.) Noot., Leiden Bot. Ser. 1:
159 (1975); S. cochinchinensis var. thwaitesii
(F.Muell.) Noot., Leiden Bot. Ser. 1: 160
(1975). Type: New South Wales. Hastings
River, s.dat ., Beckler s.n. (holo: MEL46540,
46541; iso: BRI, P).

Illustrations : Nooteboom (1982, Fig. 11)
under S. cochinchinensis var. thwaitesii ;
Floyd (1989: 397) as S. thwaitesii.

Shrub or tree to 20 m. Shoots and young
branchlets glabrous. Leaves coriaceous;
lamina oblanceolate, narrowly obovate or
elliptic, mostly 6-16 cm long, 1.5-5 cm wide,
acuminate, acute or obtuse at apex, acute or
attenuate at base, glabrous; margins serrate;
secondary veins mostly 8-11 pairs; petiole
5-15 mm long, glabrous. Inflorescence
paniculate or racemose to 7 cm, glabrous;
pedicels 0-8 mm long; bracts and bracteoles
obovate to triangular, glabrous or scarcely
ciliate, caducous, bracts 1.5-4 mm long,
bracteoles 1-2 mm long. Calyx limb 1.7-2
mm long, glabrous; lobes depressed obovate,
1.4-1.6 mm long glabrous. Petals broadly

249

Jessup, Symplocos in Australia

obovate, mostly 4.5-7 mm long, 3.5-5 mm
wide, white, glabrous. Stamens 80-100,
2-5 mm long. Disc glabrous. Style 4-5 mm
long, glabrous. Fruit ovoid to ampulliform,
6-11 mm long, 4-8 mm wide, blue, purple or
black.

Additional selected specimens examined : Queensland.

Port Curtis District: Isopod, Mt Parnassus, Oct 1976,
Hyland 4314RFK (BRI); Bulburin S.F. 67, Apr 1980,
McDonald 3220 et al. (BRI). Wide Bay District: The
Missings, S.F. 915 Tuan, Dec 1994, Forster PIF16046
et al. (BRI); S.F. 639 Wrattens, Beauty Spot 50, end of
Black Snake Road, Jan 2003, Haskard & Smyrell s.n.
(BRI [AQ772885]); c. 2 miles [3.2 km] NE Maleny,
May 1959, Thorne 21263 et al. (BRI); Mary Cairncross
Park, Sep 2004 , Russell s.n. (BRI [AQ613832]). Burnett
District: Bunya Mountains, Oct 1919, White s.n. (BRI
[AQ84418]). Moreton District: Brolga N.P, W of
Woombye, Feb 1990, Bean 1349 (BRI); Walking track
between Lawnton Road & Joyners Ridge Road, c. 1 km
from Lawton Road, Mt Glorious, Sep 2002, Phillips 783
(BRI); Mt Glorious c. 50 km W of Brisbane, Sep 1978,
Jessup 147 & Reynolds (BRI); Mt Mistake, May 1948,
Smith 3689 & Webb (BRI); Cainbable Range, Oct 1952,
Lahey s.n. (BRI [AQ478660]); Burleigh Heads N.P,
Tallebudgera, Jul 1978, McDonald 2165 et al. (BRI).
Darling Downs District: Mt Cordeaux, just below the

summit, Dec 1981, Guymer 1661 & Jessup (BRI); The
Head, Main Range N.P, Jan 2002, Forster PIF28066

6 Leiper (BRI); Border of Lamington N.P., Sarabah
Range, Oct 1969, Schodde 5576 (BRI). New South
Wales. Unumgar near base of Mt Lindsay, Nov 1944,
Jones s.n. (BRI [AQ84429]); Booyong Reserve NE of
Lismore, Nov 1982, Floyd s.n. (BRI [AQ393589]); c. 1
mile [1.6 km] S of Port Macquarie, Oct 1966, Schodde
5152 (BRI, CANB, L); Robertson Nature Reserve, Jun
1994, Kodela 315 et al. (BRI). Victoria. Spur between
two gullies entering Snowy River on W bank of Wood
Point [13 km NW of Orbost], Mar 1976, Cameron 6132a
(MEL).

Distribution and habitat: Endemic to eastern
Australia from central Queensland (Mt
Parnassus near Rockhampton) south to the
Snowy River in Victoria (Map 18). Grows
mostly in notophyll vine forest on substrates
derived primarily from basalts and granites.

Phenology: Flowers are recorded from
September to November; fruit in February.

Conservation status: Symplocos thwaitesii is
widely distributed and is not considered to be
threatened.

19. Symplocos wooroonooran Jessup nom. et
stat. nov.; S. stawellii var. montana C.T.White,
Proc. Roy. Soc. Queensland 50:82 (1939);
S. cochinchinensis var. montana (C.T.White)
Noot., Leiden Bot. Ser. 1: 160 (1975). Type:
Queensland. Cook District: Mt Bartle Frere,

7 October 1938, H.Flecker 857 (holo: BRI).

Illustrations: Cooper & Cooper (2004: 539);
Hyland et al. (2010), both under Symplocos
stawellii var. montana.

Tree to 8 m. Shoots appressed pubescent,
glabrescent; young branchlets glabrescent.
Leaves chartaceous to thinly coriaceous;
lamina oblanceolate to elliptic, 2-6.5 cm
long, 0.9-2.4 cm wide, acuminate at apex,
mostly attenuate at base, glabrous or nearly
so; margins recurved, remotely denticulate or
sometimes entire; secondary veins mostly 7-
10 pairs; petiole 2.5-4.5 mm long, glabrescent
or glabrous. Inflorescence axes 2 or 3,
racemose, to 1.5 cm long, lightly appressed
puberulous, branching at base, sometimes
single; pedicels 0-1 mm long; bracts and
bracteoles ovate, sparsely puberulous, ciliate,
mostly persistent until after anthesis, bracts 1-
2 mm long, bracteoles 0.6-1 mm long. Calyx
limb 0.75-1 mm, glabrous; lobes depressed
ovate, triangular or shortly oblong, rounded,

250

0.5-0.8 mm long, glabrous. Petals oblong
or obovate, 4-5 mm long, white, glabrous.
Stamens mostly 30-35. Disc pubescent. Style
3-3.5 mm long, pubescent near base. Fruit
ampulliform, 5.8-6.3 mm long, 4-5.5 mm
wide, black. Fig. 3E-G.

Additional selected specimens examined: Queensland.
Cook District: S.F.R. 310, Upper Goldsborough L.A.,
Sep 1976, Dockrill 1278 (BRI); Bellenden Ker, s.dat.,
Johnson s.n. (BRI [AQ519814]; MEL46558, 46559,
46560,46561,46562); Summit of Mt Bellenden Ker, Aug
1971, Hyland 5304 & 5316 (BRI); N.P.R. 226, Bellenden
Ker, Jan 1972, Hyland 5775 (BRI); Bellenden Ker
Range, Oct 1974, Hyland 7769 (BRI); Centre Peak, near
TV tower, summit of Bellenden Ker, Nov 1972, Webb &
Tracey 10802 (BRI); Mt Bellenden Ker, Jun 1969, Smith
14630 (BRI); loc. cit.. May 2001, Cooper 1525 & Cooper
(BRI); Wooroonooran N.P, Mt Bellenden Ker summit
area, Dec 2000, Forster PIF26516 et al. (BRI); Mt Bartle
Frere, Oct 1935, Blake 9843 (BRI); loc. cit., Nov 1966,
Gilmour 4076 (BRI); S.F.R. 755, Gosschalk L.A., Gray
702 (BRI); Summit area of Mt Bartle Frere, Sep 1977,
Powell 833 et al. (BRI); Wooroonooran N.P, Bartle
Frere, first heath to SE of South Peak, Oct 1997, Forster
PIF21767 et al. (BRI).

Distribution and habitat : This species is
endemic to the Wet Tropics of Queensland
where it occurs only on the upper slopes and
summits of Mt Bartle Frere and Mt Bellenden

Austrobaileya 8(3): 225-251 (2011)

Ker in Wooroonooran N.P. (Map 19). It grows
in montane rainforest (simple microphyll
vine-fern thicket and shrubland) on granite
derived substrates.

Phenology : Flowers are recorded in October
and November; fruit from January to May.

Notes : This species was treated as Symplocos
cochinchinensis var. montana (C.T.White)
Noot. (as S. cochinchinensis subsp. thwaitesii
var. montana) in Nooteboom (1982).

Conservation status : Symplocos

wooroonooran (as Symplocos stawellii
var. montana) is currently listed as Near
Threatened under the NCA.

Etymology : The species epithet

commemorates the botanically diverse
Wooroonooran N.P. and is also the aboriginal
name for Mt Bellenden Ker (Bailey 1900a).

Acknowledgements

This work was partially funded by a grant
from the Australian Biological Resources
Study (ABRS) for the purposes of providing
an account of the family for the Flora of
Australia. I would like to thank the directors
and staff of BM, BRI, CANB, CNS (formerly
QRS), K, L, MEL, NSW and P for access to
their collections. Will Smith produced the
line drawings and Peter Bostock checked and
corrected my Latin diagnoses.

References

Bailey, F.M. (1890). A Synopsis of the Queensland
Flora, Supplement 3: 46. H.J.Diddams & Co.:
government Printer: Brisbane.

- (1900a). Leptospermum wooroonooran

F.M.Bailey. In The Queensland Flora 2: 588.
H.J.Diddams & Co.: Government Printer:
Brisbane.

- (1900b). Symplocos Linn. In The Queensland

Flora 3: 967-968. H.J.Diddams & Co.:
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Bentham, G. (1868). Symplocos Linn. (Styracaceae). In
Flora Australiensis 4: 292-293. L. Reeve & Co.:
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Brand, A. (1901). Symplocaceae. In H.G.A.Engler (ed.),
Das Pflanzenreich Heft 6(1 V, 242): 1-111.
Engelmann: Leipzig.

Map 19. Distribution of Symplocos wooroonooran

251

Jessup, Symplocos in Australia

Bostock, P.D. & Holland, A.E. (eds.) (2010). Census
of the Queensland Flora 2010. Queensland
Herbarium, Department of Environment and
Resource Management: Brisbane.

Cooper, W. & Cooper, W.T. (2004). Fruits of the
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Floyd, A.G. (1989). Rainforest Trees of Mainland South¬
eastern Australia. Inkata Press: Melbourne/
Sydney.

Fritsch, P.W., Cruz, B.C., Almeda, F., Wang, Y. & Shi,

5. (2006). Phylogeny of Symplocos based on
DNA sequences of the chloroplast trnC-trnD
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192.

Fritsch, P.W., Kelly, L.M., Wang, Y., Almeda, F. &
Kriebel, R. (2008). Revised infrafamilial
classification of Symplocaceae based on
phylogenetic data from DNA sequences and
morphology. Taxon 57: 823-852.

Green, P.S. (1994). Symplocaceae. In Flora of Australia
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Hyland, B.P.M., Whiffin, T. & Zich, F. (2010).
Australian Tropical Rainforest Plants. Edition

6. http://kevs.trin.org.au:8080/kev-server/data/

0e0f0504-0103-430d-8004-060d07080d04/

media/Html/index.html

Jessup, L.W. (1993). Two new species of Symplocos
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- (1994). Symplocaceae. In R.J.F. Henderson

(ed.), Queensland Vascular Plants: Names and
Distribution , p. 329. Queensland Herbarium:
Indooroopilly.

- (1997). Symplocaceae. InR. J.F. Henderson (ed.),

Queensland Plants: Names and Distribution ,
p. 201. Queensland Herbarium, Department of
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Leiper, G., Glazebrook, J., Cox, D. & Rathie, K. (2008).
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Australian Plants (QLD Region) Inc.: Browns
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Mueller, F. (1862). FragmentaPhytographiaeAustraliae
3: 22. Government Printer: Melbourne.

- (1865). Fragmenta Phytographiae Australiae 5:

60. Government Printer: Melbourne.

Nooteboom, H.P (1975). Revision of the Symplocaceae
of the Old World, New Caledonia excepted.
Leiden Botanical Series 1:16+336. Universitaire
Pers Leiden: Leiden.

- (1982). A revision of the Australian species of

Symplocos (Symplocaceae), Brunonia 4: 309-
326.

Smith, A.C. (1978). A precursor to a new Flora of Fiji.
Allertonia 1: 375-377.

- (1981). Symplocaceae. In Flora Vitiensis Nova

2: 724-729. Pacific Tropical Botanical Garden:
Lawai, Kauai, Hawaii.

Stanley, T.D. & Ross, E.M. (1986). Symplocaceae. In
Flora of south-eastern Queensland 2: 272-273.
Queensland Department of Primary Industries:
Brisbane.

Thomas, M.B. & McDonald, W.J.F. (1989). Rare and
Threatend Plants of Queensland. 2nd Edition.
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Brisbane.

Wang, Y., Fritsch, P.W., Shi, S., Almeda, F., Cruz, B.C. &
Kelly, L.M. (2004). Phylogeny and infrageneric
classification of Symplocos (Symplocaceae)
inferred from DNA sequence data. American
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- (2002). Symplocaceae. In R.J.F. Henderson

(ed.), Names and Distribution of Queensland
Plants, Algae and Lichens , p. 193. Queensland
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- (2007). Symplocaceae. In P.D. Bostock & A.E.

Holland (eds.). Census of the Queensland Flora
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Management: Brisbane.

A taxonomic revision of Pseudoweinmannia
Engl. (Cunoniaceae: Geissoieae)

Andrew C. Rozefelds 1 & Belinda Pellow 2

Summary

Rozefelds, AC. & Pellow, B. (2011). A taxonomic revision of Pseudoweinmannia Engl. (Cunoniaceae:
Geissoieae). Austrobaileya 8(3): 252-266. The vegetative and reproductive morphology of
Pseudoweinmannia is described and illustrated. The genus is endemic to eastern Australia with two
species, Pseudoweinmannia apetala (F.M.Bailey) Engl, and P. lachnocarpa (F.Muell.) Engl., that
can be differentiated by their leaf serration, stipule morphology and flower and fruit characters.
New interpretations of floral and fruit morphology in Pseudoweinmania are discussed, and their
significance in better understanding the phylogenetic relationships within the tribe Geissoieae are
examined.

Key Words: Cunoniaceae, Geissoieae , Pseudoweinmannia , Pseudoweinmannia apetala ,
Pseudoweinmannia lachnocarpa , Australia flora, Queensland flora. New South Wales flora,
comparative morphology

‘Andrew C. Rozefelds, Queensland Museum, GPO Box 3300, South Brisbane, Queensland 4101,
Australia. Email: andrew.rozefelds@qm.qld.gov.au

2 Belinda Pellow, Janet Cosh Herbarium, Department of Biological Sciences, University of
Wollongong, Northfields Avenue, Wollongong, New South Wales 2522, Australia.

Introduction

The family Cunoniaceae includes 15 genera
in Australia, which are all primarily rainforest
trees and/or shrubs, and are currently being
revised for the Flora of Australia. Recent
revisions have been completed on the genera:
Anodopetalum A.Cunn ex Endl. (Barnes
& Rozefelds 2000), Ceratopetalum Sm.
(Rozefelds & Barnes 2002), Davidsonia
F.Muell. (Harden & Williams 2000), the
Australian species of Geissois Labill.
(Schimanski & Rozefelds 2002), Vesselowskya
Pamp. (Rozefelds et al. 2001) and a new
species of Gillbeea F.Muell. was described
from north eastern Australia (Rozefelds &
Pellow 2001). In this paper a revision of the
Australian endemic genus Pseudoweinmannia
Engl, is presented.

Pseudoweinmannia was proposed by
Engler (1930) for two species, Weinmannia
lachnocarpa F.Muell. and W. apetala
F.M.Bailey from eastern Australia, which he
referred to the tribe Cunonieae (R.Br.) Schrank
& Mart, in Cunoniaceae. Pseudoweinmannia
differs from Weinmannia, the major genus

Accepted for publication 8 August 2011

in this tribe, in lacking petals, having a
polyandrousandroeciumandtardily-dehiscent
fruits. Based upon an analysis of stipule
morphology Dickison & Rutishauser (1990)
concluded, that Pseudoweinmannia was more
closely related to Geissois and Lamanonia
Veil, in the Tribe Belangerae Lindl ex Pfeiff.
(= Tribe Geissoieae Endl. ex Meisn.), than to
the Tribe Cunonieae.

In a cladistic analysis using morphological
characters, Hufford & Dickison (1992) showed
that Pseudoweinmannia was sister to Geissois
(including Lamanonia ), as it shared with this
genus, lateral stipules (Dickison & Rutishauser
1990), racemose inflorescences and a
polyandrous androecium. Dickison (1984)
and Hufford & Dickison (1992) also identified
structures on the seeds as elaiosomes, which
they interpreted as a putative autapomorphy
for the genus.

Combined morphological and molecular
data based upon analysis ofrZ>cL, and intron and
spacer regions in the combined trnL-G region,
and chloroplast DNA data provide support for
interpreting Pseudoweinmannia and Geissois
sens. lat. as sister taxa (Bradford & Barnes
2001). These authors recircumscribed the

Rozefelds & Pellow, Revision of Pseudoweinmannia

253

Tribe Geissoieae (= Tribe Belangerae sensu
Dickison & Rutishauser 1990) to include the
Australian and New Caledonian species of
Geissois , Lamanonia from South America,
and Pseudoweinmannia.

Pseudoweinmannia is endemic to eastern
Australia where it is restricted to rainforests.
Pseudoweinmannia lachnocarpa (F.Muell.)
Engl. (Fig. 1) occurs in north eastern New
South Wales and south eastern Queensland
(Map 1), and was described by Mueller (1874)
from material collected from the Tweed
District in north eastern New South Wales. A
second species, Pseudoweinmannia apetala
(F.M.Bailey) Engl. (Fig. 2), was described by
Bailey (1893), from Kamerunga, near Cairns
in north eastern Queensland (Map 1). Bailey
(1893), in describing Pseudoweinmannia
apetala , noted that it “approaches near to
lachnocarpa but differs in its more lax slender
inflorescence, more membranous leaves,
which are not so prominently reticulate, its
longer stamens, as well as probably the flower
lobes to the calyx”. Since their description
there has not been a critical re-examination of
the morphology of both species and as a result
the status of Pseudoweinmannia apetala has
become unclear. Recent reviews in flora and
state censuses have either considered it a
synonym of Pseudoweinmannia lachnocarpa
(e.g. Henderson 1997; Hyland & Whiffin
1993; Jessup 2010), or a distinct species (e.g.
Stanley & Ross 1983; Harden 1990; Bradford
etal. 2004).

In this paper the vegetative and
reproductive morphology of the two species
is described using herbarium collections
(both dried and in spirit) and consistent
morphological differences are recognized
between Pseudoweinmannia lachnocarpa and
P. apetala. The vegetative and reproductive
morphology of some key characters in
Pseudoweinmannia, i.e. stipule morphology,
inflorescence architecture and floral and fruit
morphology, are also described and illustrated
to permit comparisons with other genera in
the Tribe.

Methods and materials

Herbarium collections examined include those
from BRI, CANB, CNS (formerly in QRS),
K, NSW and MEL. The principal collections
used are cited herein and specimens examined
with the scanning electron microscopy (SEM)
are indicated by *. Common abbreviations
used in specimen citation are N.R (National
Park), L A. (Logging Area), S.F./S.F.R. (State
Forest/State Forest Reserve).

Terminology follows Walsh & Entwisle
(1999) for leaf morphology and Hickey (1979)
for leaf architecture.

For SEM the material prepared included
floral organs, pollen, seeds, and stipules. The
specimens were either air-dried or critical
point dried, and placed onto aluminium stubs
with tape. They were then sputter coated with
gold and examined with a Philips Electroscan
Environmental Scanning Electron Microscope
2020, under high vacuum operated at 10-15
kV.

Results

Stipule morphology

The stipules in Pseudoweinmannia are keel¬
shaped, lanceolate structures. Developmental
studies in Pseudoweinmannia and the
Australian Geissois species by Dickison
& Rutishauser (1990) showed that four
stipular primordia arise laterally at each
node. In Pseudoweinmannia the two stipular
primordial, on either side of the node, coalesce
to form a bifid laterally-positioned compound
structure (Fig. 3B). The margins of the
stipules have glandular trichomes and simple
hairs in Pseudoweinmannia apetala while in
P. lachnocarpa only glandular trichomes are
present (Fig. 3A-C).

Inflorescence architecture

Schimanski & Rozefelds (2002) have
described the inflorescence morphology in
the Australian species of the related genus
Geissois. Their terminology is applied here
to describe the inflorescence morphology of
Pseudoweinmannia.

The flowers of Pseudoweinmannia are
arranged in indeterminate racemose units.

254

Austrobaileya 8(3): 252-266 (2011)

20 DEC 2001

Depth m
Alt. 960m

173 Cunoniaceae

AQ 567426

QUEENSLAND HERBARIUM (BRI)

Flora of Queensland Cook

Pseudoweinmannia lachnocarpa (F.Muell.) Engl.

QUEENSLAND HERBARIUM (BRI)
Brisbane Australia

557426

Coll. A, Ford
Jensen,R.;

<16d 17m 36s 145d 5m 18s)[AGD66]

(55,295750,8197650) 7965-957976
SFR144, Chovvchilla Logging Area, along old logging track, Windsor
Tableland.

Complex notophyll rainforest on soil derived from granite. Agathis
robusta very common.

Canopy of Aleurites, Argyrodendron poly end rum and
Pseudoweinmannia.

Cdfnmon small trees - Austromyrtus spp., Dinosperma, Croton
insularis and Elaeodendron.

Canopy tree 15m high. New growth reddish.

Fruit slightly immature, hairs creamish when fresh.

Dup. NSW MEL HO\LN^

* May be cited as computerised collection Number
(Archival Paper)

Fig. 1 . Representative specimen of Pseudoweinmannia apetala {Ford AF3202 & Jensen [BRI])

Rozefelds & Pellow, Revision of Pseudoweinmannia

255

QUEENSLAND HERBARIUM (BRI)
Flora of Queensland

Pseudoweinmannia lachnocarpa (F.Muell.) Engl.

Coll, P.l.Forster PIF9353
+P.Machim

Moreton

6 JAN 1992

27 b 25'S 152*49'E Alt> m .

2l State Forest 309 Enoggera. " Grid ref!

9443-o24o60.

creek 3 '" 13 ' 1 not ° phyl 1 vine forest on dark brown soil along
Tree to 5m, cream flowers.

Occasional In area.

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

AQ

520250

Fig. 2. Representative specimen of Pseudoweinmannia lachnocarpa {Forster 9353 & Machin [BRI])

256

Following Bradford (1998), we use the term
raceme to describe these elements, although
occasionally the racemes are weakly branched
(e.g. Webb & Tracey 11246). Bracts subtend
each flower, but are quickly deciduous. The
pedicels are shortly hirsute and have an
abscission point about 1-2 mm above the
point of attachment.

Variation in Psendoweinmannia , as in
the Australian species of Geissois , occurs
at the following hierarchical levels: (1) in
the varying development of the racemes
and supporting axes and buds that form the
inflorescence (Inflorescence Module sensu
Bradford 1998); and (2) in the position that
the inflorescences occupy in relation to the
main stem axis. In both Pseudoweinmannia
apetala and P. lachnocarpa the common
condition, particularly in younger shoots, is
for the raceme to occupy an axial position in
the leaf axils and a terminal bud is present.

The pattern is more complicated in
older branch units, where inflorescence
units may develop in a number of leaf axils,
and a compound inflorescence is formed;
subtending leaves, may or may not, be
present. The inflorescence could be described
as a compound homothetic inflorescence
{sensu Weberling 1989), and the arrangement
of raceme-like units on the flowering stem is
consistent with the decussate architecture of
the shoot. In some specimens the terminal
bud may also be replaced by a raceme
{Pseudoweinmannia lachnocarpa {Blake
22848)' P. apetala {Whiffin 5978 , Gray
5978)). Heterothetic compound inflorescences
{sensu Weberling 1989) therefore result if the
terminal bud is replaced by a raceme-like unit
and inflorescences are initiated in a number
of leaf axils. In both Pseudoweinmannia
apetala and P. lachnocarpa , heterothetic and
homothetic compound inflorescences may
occur on the same specimen, and no consistent
inflorescence architectural differences have
been identified between the two species.
The racemes are somewhat variable in
length, although those of Pseudoweinmannia
apetala are usually longer than that in
P. lachnocarpa.

Austrobaileya 8(3): 252-266 (2011)

Floral and fruit morphology

TheflowersinbothPsezzdowzzzzzzawzzaspecies
are (5-)6(-7)-merous and polystemonous
(Bradford & Barnes 2001; Bradford etal. 2004;
Schimanski & Rozefelds 2002) (Fig. 4B,E).
The sepals are valvate, ovate in shape and have
a strigose indumentum over their inner and
outer surfaces (Fig. 4A,D). Petals are lacking
in both species (Fig. 4B,E). Anthers are
introrse, dorsifixed at just below the midpoint
of the anther and are weakly versatile and
lack a sterile distal protrusion (Fig. 4A,D,E).
A prominent, lobed annular nectary occurs at
the base of the ovary (Fig. 4B,F). The ovary is
bicarpellate, rarely tricarpellate, syncarpous,
and is covered by a dense indumentum of
strigose hairs (Fig. 4A,B>C,F). The styles
are furrowed ventrally, are slightly coiled
in bud and in young flowers (Fig. 4A) and
straighten with flower development (Fig. 4B).
The styles are exserted before the anthers
and while studies on stigmatic receptivity
have not been undertaken, the flowers are
possibly proterogynous (Fig. 4D). The stigma
is papillate at maturity (Fig. 4C). The fruit
is a tardily dehiscent capsule. The ovary
in Pseudoweinmannia is divided into two
parts, rarely three, indicating origin from
two carpels; the two carpels are adnate and
free (Fig. 5A,B) The seeds are irregularly
rounded pyramidal in shape (Fig. 6A,C), with
weakly reticulate cell walls (Fig. 6B,D) and
lack a wing-like extension.

Comparative morphology in the Tribe

Geissoieae

The Tribe Geissoieae consists of
Pseudoweinmannia , Lamanonia and Geissois
sens, lat., and includes species that occur in
Australia, South America and New Caledonia
(Bradford & Barnes 2001). The adult leaves
in Pseudoweinmannia are trifoliolate and
have two caducous stipels at the base of the
lateral leaflets. Australian Geissois species
also have stipels, or rarely reduced leaflets,
that occupy the same position as those in
Pseudoweinmannia (Schimanski & Rozefelds
2002). The New Caledonian Geissois taxa
and Lamanonia are typically digitate or
trifoliolate, the most basal leaflets being
the smallest. Bradford & Barnes (2001)

Rozefelds & Pellow, Revision of Pseudoweinmannia

257

Fig. 3. Variation in stipule morphology in Pseudoweinmannia apetala and P. lachnocarpa. A. P. lachnocarpa,
detail of stipule margin showing glandular trichomes. B. P. apetala, the pair of stipules and the margins between the
two stipules are arrowed. C. P. apetala stipule, detail of stipule margin showing unicellular and glandular trichomes
(arrowed). Scale bars A,C = 300 pm; B = 1 mm. A from Floyd s.n. (CANB 289092); B-C from Gray 1272 (CNS).

interpreted the common leaf condition in the
tribe as being digitate (palmately compound),
which is logical if the stipels in Geissois
and Pseudoweinmannia are interpreted as
homologues to lateral leaflets.

Dickison (1975) described the stomata in
Pseudoweinmannia as anomocytic, although
recent studies by Barnes in Bradford &
Barnes (2001) interpreted the stomata as
brachyparacytic, which is the common
condition in the family and is present for all
genera of the Tribe Geissoieae. The secondary
leaf venation is semi-craspedodromous,
which is the common condition in most of
the Cunoniaceae and is typical of Geissoieae
(Hufford & Dickison 1992; Barnes pers.
comm. 2011).

The androecium is defined as polyandrous
in the Geissoieae (Bradford & Barnes 2001).
In the New Caledonian species of Geissois ,
developmental studies show that the stamens
are grouped together in fascicles, occupy

an episepalous position and development
proceeds from a centre of the fascicle
outwards in both directions (Matthews &
Endress 2002). In developing flowers the
inner stamens are typically larger than the
outer ones in the fascicle. This arrangement
of stamens is evident in older stages of flower
development in the Australian species of
Pseudoweinmannia (Rozefelds pers. obs .)
and Geissois (Schimanski & Rozefelds 2002,
figs 25, 27). The arrangement of stamens
in Pseudoweinmannia and the Australian
species of Geissois is considered to be weakly
polyandrous.

The common fruit type in the Cunoniaceae
is a bilocular, ventrally dehiscent capsule
(Dickison 1984). Geissois s.l. and Lamanonia
have ventrally dehiscent capsules, while those
of Pseudoweinmannia were interpreted by
Dickison (1984) as an ‘indehiscent capsule’
and Bradford et al. (2004) similarly referred to
the fruits in Pseudoweinmannia as indehiscent

258

Austrobaileya 8(3): 252-266 (2011)

Rozefelds & Pellow, Revision of Pseudoweinmannia

fruits. Other researchers have indicated that
the fruits are septicidally dehiscent (Harden
1990) and open into two valves (Floyd 1989).
The semantic discontinuities imposed by
terms, such as ‘indehiscent’ versus ‘dehiscent’,
obscure the morphological similarities that
link these genera together.

The fruits in Pseudoweinmannia are
typically derived from two carpels, although
they are obscured by the mass of fine hairs
on the outside of the fruit (Dickison 1984,
fig. 13). Sections through the fruit indicate
that the walls of the carpels are adnate, but
are not fused together (Fig. 5A,B) The
placental proliferations that Dickison (1984,
pi. 2, Fig. 21) illustrated were not evident in
the specimens we sectioned. The thickened
(sclerenchyma) walls of the capsules can
be convoluted in appearance as the oblique
section revealed (Fig. 5B), but near the mid¬
region of the fruit these walls are straight
(Fig. 5B). It seems likely that Dickison’s
interpretations were based upon either a
section through an immature fruit and/or
an oblique section through the fruit that cut
through the developing, convoluted walls of
the capsule.

InPseudoweinmanniathQ fruits are similar
to those of the other genera as the walls of the
two carpels are not fused together but stay
conjoined for some time after fruit fall. The
mid-section through the Pseudoweinmannia
fruits indicates that sclerenchyma occupies
a largely identical position to that seen in
Geissois stipularis A.C.Sm (Dickison 1984, pi

Fig. 4. (facing page) Floral morphology of
Pseudoweinmannia apetala and P. laclinocarpa. A-C.
P. apetala. A. Partially opened bud with some stamens
removed to show dense covering of hairs on ovary, and
curved styles. B. Mature flower with fully exserted
stamens, and the two style branches are arrowed, note
conspicuous disc (nectary = N) at base of ovary. C. Top
view of flower and the papillate surface of the stigma
is arrowed (white arrow). D-F. P. lachnocarpa. D.
partially opened bud, note that the style branches are
exserted (white arrows) and the anthers are dorsifixed
at just beiow anther midpoint (black arrow). E. Newly
open flower, note the variation in length of the stamen
filaments. F. Mature flower with stamen filaments
extended and basal disc (N). Scale bars A,C,D = 1 mm;
B,E,F = 2 mm. A from Gray 5978 (CNS); D-E from
Rozefelds s.n. (HO); F from Forster PIF9353 & Machin

259

2, fig. 22). All genera in the Tribe Geissoieae
therefore have bi-locular fruits that are
derived from capsules and are dehiscent or
tardily dehiscent.

Pseudoweinmannia differs from the other
genera in the Tribe by stipule morphology
(Schimanski & Rozefelds 2002) and in
seed and fruit morphology. The stipules
in Pseudoweinmannia are keel-shaped,
lanceolate structures (Fig. 3B), while those
in Geissois from Australia (Schimanski &
Rozefelds 2002) and Lamanonia from South
America are large and often foliose (Leite
1983; Zickel & Leitao Filho 1993; Engler
1930).

Pseudoweinmannia has rounded,
pyramidal seeds while the other genera in
the tribe have winged seeds. Dickison (1984)
described the seeds of Geissois s. lat. as
having distal wings. Lamanonia also has a
distal wing (Engler’s (1930) illustration of
L. tomentosa Camb. (= 1L. ternata Veil.)',
Zickel & Leitao Filho 1993; Leite 1983),
and Dickison (1984) interpreted the seed in
Lamanonia ternata as having a distal wing
and lateral extensions. Dickison (1984) drew
particular attention to an aril-like outgrowth
on the seeds in Pseudoweinmannia , which
he interpreted as an elaiosome. We observed
a few, thin-walled parenchyma cells around
the point of attachment of the seed, but we
interpret these cells as being derived from the
placental parenchyma (Fig. 5B,C). We are
unaware of any ecological or field studies that
support interpretation of these structures as
an elaiosome.

The seeds of Pseudoweinmannia were
described as having either a smooth (Dickison
1984) or papillate (Hufford & Dickison 1992)
surface, although the material we examined of
both species had a smooth, weakly reticulate
appearance due to the cell walls (Fig. 6A-D).
Dickison (1984) considered the epidermal
cells of the Australian species of Geissois to be
more ornamented and they could be identified
apart by differences in the epidermal cell
structures. The epidermal cells of the seed
coats of Lamanonia and Geissois stipularis
A.C.Sm. (New Caledonia) were described
as having irregular outlines and moderately
thick to thin lateral walls (Dickison 1984).

260

Austrobaileya 8(3): 252-266 (2011)

Fig. 5. A. Pseudoweinmamtia lachnocarpa, oblique
transverse section through a mature fruit. Note the
position of the seeds in the fruit and the thickened seed
coat (SC). B. Drawing of A showing that the walls of
the two carpels are adnate but free; the sclerenchyma
wall (sc) of the right hand carpel are not continuous and
remain free near the mid-section of the fruit. The more
oblique section through the left hand carpel shows that
the convoluted sclerenchyma (sc) wall of the carpel is
continuous. C. Simplified transverse section of fruit of
Geissois stipularis A.C.Sm., redrawn from Dickison
(1984), note that the sclerenchyma walls (sc) of the
capsule remain free. (S = seed; SC = seed coat; sc =
sclerenchyna). Scale bar = 1 mm. A from Williams s.n.
(MEL).

The tribe Geissoieae are a very cohesive
group, with all species having a similar
vegetative and reproductive morphology. The
species share digitate (palmately compound)
leaves, brachyparacytic stomates and semi-
craspedodromous secondary venation. The
floral plan in all genera consists of valvate
sepals, no petals, polyandrous androecium,
the anthers lack a distal connective protrusion,
and the bicarpellate ovary is covered in a thick
indumentum with a prominent disc (nectary)
at the base. Pseudoweinmannia also shares
with the other two genera dicolporate pollen,
raceme-like inflorescences and capsular
dehiscent fruits (Hufford & Dickison 1992;
Bradford & Barnes 2001; Schimanski &
Rozefelds 2002; herein).

Systematics

Family Cunoniaceae

Tribe Geissoieae Endl. ex Meisn.

Pseudoweinmannia Engl., Nat. Pflanzenfam.
2nd Ed., Vol. 18a: 249 (1930). Type species:
Pseudoweinmannia lachnocarpa (F.Muell.)
Engl. (= Weinmannia lachnocarpa F.Muell.)

Trees or shrubs. Leaves opposite, decussate,
trifoliolate and with a pair of stipels at base
of lateral leaflets (digitate); unifoliolate in
coppice growth or seedlings; leaflets sessile
or very shortly petiolulate, similar in size,
central leaflet sometimes up to one-third
larger, broadly lanceolate, elliptical to obovate,
apices typically acute, less commonly obtuse,
leaf bases are cuneate to attenuate, lateral
leaflets base slightly asymmetrical, margin
serrate to crenate, sinuses weakly to strongly

Rozefelds & Pellow, Revision of Pseudoweinmannia

developed, venation semi-craspedodromous,
midrib course straight in central leaflet and
slightly curved in lateral leaflets, secondary
venation thickness stout, alternate, uniformly
divergent, tertiary venation percurrent,
complete areolation is formed by higher-
order venation, teeth dark, glandular with
simple apical termination. Stipules 4 at each
node, keel-shaped, lanceolate, apex acute.
Inflorescences usually axillary, consisting of
raceme-like units, which typically occur in the
upper leaf axils, or occasionally terminate the
main axis. Flowers bisexual; sepals (5-)6(-7)
free; petals absent; androecium weakly

261

polystemonous, stamens 12-25, carpels 2 (or
3) fused, styles subulate, 2 (or 3) distally free,
ovary densely hairy, stigmas papillate, 2-4
ovules/carpel, disc (annular nectary) around
base of ovary. Fruit a dry, tardily dehiscent
capsule covered in a dense, woolly golden-
brown indumentum, stamen filaments and
styles persistent; carpels free; seed rounded-
pyramidal, not winged, seed walls weakly
reticulate.

Etymology: Name from the Latin pseudo
meaning (false), and Weinmannia a genus
named in honour of Johann Wilhelm
Weinmann, German pharmacist and botanist,
(1683-1741).

Key to Pseudoweinmannia species

Leaf margins crenulate-crenate, sinuses weakly developed; stipules
fringed with glandular and simple trichomes; stamens

12-20, disc < 2.5 mm high; fruit ovoid-globose. NE Qld. P. apetala

Leaf margins serrulate-serrate (rarely crenate), sinuses well developed;
stipules fringed with glandular trichomes; stamens (12-)20-24, disc
>3 mm high; fruit globose. SE Qld, NE NSW. P. lachnocarpa

Pseudoweinmannia apetala (F.M. Bailey)
Engl, in Engl. & Prantl., Nat. Pflanzenfam.
2nd Ed., Vol. 18a: 249 (1930); Weinmannia
apetala F.M.Bailey, Dept. Agric. Brisbane,
Bot. Bull. 8: 76 (1893). Type: Queensland.
Cook District: Kamerunga, January 1892,
E.Cowley 65, ‘the growing shoots from roots,
flower’ (holo: BRI [AQ341025]; iso: K n.v.
[photo at HO!]).

Pseudoweinmannia lachnocarpa auct. non
(F.Muell.) Engl.; Hyland & Whiffin (1993:
131); Cooper & Cooper (2004: 148).

Illustrations: Christophel & Hyland (1993:
pi. 25, fig. C); Cooper & Cooper (2004: 148);
all as P. lachnocarpa.

Tree up to 30 m tall, often prominently
buttressed in larger trees (Hyland & Whiffin
1993). Outer bark flaky ( Sanderson 1634).
Branchlets terete, bark grey with conspicuous
lenticels. Stipules 2-5 mm long, base to 2 mm
wide, margin entire with glandular trichomes
(colleters) and non-glandular strigose
trichomes, glabrous above, caducous. Leaves
opposite, decussate, petiolate, trifoliolate with

two caducous stipels; petiole 1.5-4.5 cm long,
broadly elliptic in cross-section, glabrous to
sparsely strigose; leaflets 5-12.5 cm long,
1.5-4.9 cm wide, sessile or occasionally with
short petiolule, broadly lanceolate, elliptical
to obovate, apex acute to acuminate, rarely
obtuse, bases cuneate to attenuate, margins
typically crenulate to crenate, sinuses weakly
developed; secondary veins 10-16 pairs,
60-80° to midrib and branching well inside
margin. Inflorescence usually axillary,
consisting of a simple raceme, 4.5-12 cm
long, occasionally with one or two racemose
lateral branches, or with some of the lower
flowers replaced by 3-flowered dichasia;
rachis terete, shortly hirsute with crispate
trichomes. Flowers bisexual; pedicels terete,
3-8 mm long, shortly hirsute with crispate
trichomes, abscission point 0.3-0.9 mm above
attachment; sepals valvate in bud, (5-)6(-7),
ovate, elliptical to narrowly ovate, 1.5-2.5
mm long, 0.6—1 mm wide, apex acute, densely
hirsute distally, trichomes more sparsely
distributed below; stamens 12-20; filaments
of unequal length, 2-4 mm long, glabrous,
anthers introrse, 0.4-0.5 mm long; disc

262

Austrobaileya 8(3): 252-266 (2011)

Fig. 6. Seed morphology in Pseudoweinmannia apetala andP. lachnocarpa. A & B. P. lachnocarpa. A. Seed showing
rounded, pyramidal shape and irregular surface and angular margins. B. Detail showing reticulate appearance of the
cell walls. C & D. P. apetala. C. seed showing rounded, pyramidal shape and angular margins. D. Detail showing
somewhat more wrinkled, reticulate appearance of the cell walls. Scale bars A = 1 mm; B,D = 200 pm; C = 0.5 mm. A
& B from Webb & Tracey 11246 (CANB); C&D from Risley 35 (NSW).

colour not noted, c. 0.2 mm high, glabrous,
slightly lobed; ovary subglobular-conical,
0.6-1 mm high, 0.9-1 mm wide, externally
woolly hirsute, 2 (or 3) celled, each cell with
two rows of 3 or 4 ovules, tapering into 2 (or
3) free styles; styles subulate, 1.8-2.8 mm
long, c. 0.2 mm wide at base, glabrous except
for a few trichomes near the base. Fruit ovoid-
globose, 3-4.5 mm high, 4-5 mm wide; seed
brown. Fig. 1.

Additional selected specimens examined : Queensland.

Cook District: T.R. 176, Monkhouse, Oct 1982, Hyland
12175 (CNS); Shiptons Flat on Tin Mine Road, May
1969, Smith 14341 (BRI); E.P 40, S.F.R. 144, Jun 1979,
Sanderson 1634 (CNS); S.F.R. 144, Dagmar, Chowchilla
L.A.,Feb 1995, Gray5978* (CNS); S.F.R. 144 Chowchilla
L.A., Windsor Tableland, Dec 2001, Ford AF3202 &

Jensen (BRI, CNS, HD); S.F.R. 1073, Dulanban, Hartley
L.A., Feb 1993, Hyland 14641* (CNS); near Kuranda,
Jan 1972, Hyland 5795 * (BRI, CNS); S.F.R. 933, Little
Pine L.A., Feb 1979, Gray 1272 (BRI, CNS); S.F.R. 185,
Downfall L.A., Mar 1973, Whiffin & Risley 586 (CNS);
S.F.R. 185, Downfall L.A., Jan 1985, Gray 3848* (CNS);
Downfall L.A., S.F.R. 185, Feb 1972, Risley 35* (BRI,
CANB, CNS, NSW); Atherton District, Dansie 2053
(BRI); Bellenden Ker Range near Babinda, Jan 1979,
Jago 270* (CNS); Downfall Creek Road, near Tinaroo
Dam, s.dat., Webb & Tracey 7646 (BRI); Babinda, May
1997, Jago 634 (CNS). North Kennedy District: Tully
River, rafting access point no. 9, SE of Cardstone, Jun
2001, FordAF2892 (BRI, CNS).

Distribution and habitat: Restricted to north
eastern Queensland from near Cardstone in
the south to Rossville near Cooktown in the
north (Map 1). Hyland & Whiffin (1993)

Rozefelds & Pellow, Revision of Pseudoweinmannia

noted that Pseudoweinmannia apetala (under
the name P. lachnocarpa) occurs from sea
level to 800 m, commonly in drier rain forest,
but also occasionally in wetter rainforest
communities. Cooper & Cooper (2004)
extended the altitudinal range to 950 m.

Phenology: Pseudoweinmannia apetala

flowers from January through to April, rarely
as late as June; with fruiting specimens
collected from December through to March.

Notes : The notes with the type specimen
i.e. “the growing shoots from roots, flower”
suggest that it may have been collected from
coppice growth at the base of the tree.

Etymology: The species name, apetala , comes
from the Greek a , not, and Latin petalum for
petals, which refers to the lack of petals in the
flowers.

Although Cooper & Cooper (2004)
list a common name as ‘Marara’, this
application is derived from populations of
Pseudoweinmannia lachnopetala and the
adoption of this aboriginal name by people
in southern Queensland and New South
Wales (Bailey 1900, 1913; Maiden 1922). No
common names were listed for this species by
Bailey (1900, 1913).

Pseudoweinmannia lachnocarpa (F.Muell.)
Engl, in Engl. & Prantl., Nat. Pflanzenfam.
2nd Ed., Vol. 18a: 249 (1930); Weinmannia
lachnocarpa F.Muell., Fragm. 8: 7 (1874);
Windmannia lachnocarpa (F.Muell.) Kuntze,
Revis. Gen. PI. 1: 228 (1891); Geissois
lachnocarpa (F.Muell.) Maiden, Forest FI.
New South Wales 7:6 (1922). Type: New South
Wales. Tweed, s.dat., [W]Guilfoyle s.n. (lecto
[here designated]: MEL104457; isolecto: K
n.v., photo at HO!; Tweed, s.dat., C.Moore s.n.
(lectopara: MEL104455, MEL104456).

Weinmannia lachnocarpa var. parvifolia
F.M.Bailey, Queensland Agric. J. 28:
196 (1912). Type: Queensland. Moreton
District: Macpherson Range, February 1912,
C.T.White s.n. (holo: BRI [AQ321400]; iso:
NSW104762).

Illustrations: Maiden (1922, t. 229); Francis
(1951: figs 11, 87, 88); Beadle (1976: fig. 150C);
Stanley & Ross (1983: Fig. 36D1,D2); Harden
(1990: 518).

263

Tree up to 40 m tall, often prominently
buttressed in larger trees (Francis 1951;
Beadle 1976). Outer bark grey or fawn pink
usually rough with small scales and pustules
(Floyd 1989). Branchlets terete, bark grey
with conspicuous lenticels. Stipules 2-3.5
mm long, base to 1 mm wide, margin entire
with glandular trichomes (colleters), glabrous
above, caducous. Leaves opposite, decussate,
petiolate, trifoliolate with two caducous
stipels; petiole 1.5-3.5 cm long, broadly
elliptic in cross-section, glabrescent, leaflets
2-13.3 cm long, 0.5-5.8 cm wide, sessile or
occasionally with a short petiolule, broadly
lanceolate, elliptical to obovate, apex acute
to obtuse and ± acuminate, base cuneate to
attenuate, margins serrulate to serrate, rarely
crenate, sinuses typically strongly developed;
secondary veins 11—16(—20) pairs, at 60-80°
to midrib and branching well inside margin.
Inflorescence typically consisting of a simple
raceme, 3-10 cm long, occasionally with one
or two racemose lateral branches, or with some
of the lower flowers replaced by 3-flowered
dichasia, rachis terete, shortly hirsute, with
crispate trichomes. Flowers bisexual; pedicels
terete, 3-8 mm long, with shortly hirsute
crispate trichomes on rachis, abscission point
0.3-1 mm above attachment; sepals valvate
in bud, (5-)6(-7), ovate-elliptical to narrowly
ovate, 2-3 mm long, 0.9-1.2 mm wide, apex
acute, densely hirsute distally and proximally,
with sparse glands on margins; stamens
(12-)20-24; filaments of unequal length 3-5
mm long, glabrous, anthers introrse, 0.5-0.8
mm long; disc red-brown in colour, c. 0.4 mm
high, distinctly lobed; ovary globular, 0.8-1
mm high, 0.9-1 mm wide, externally woolly
hirsute, 2(-3) celled, each cell with two rows
of 2, 3 or 4 ovules, tapering into 2(-3) free
styles, styles subulate, 2-3 mm long, c. 0.35
mm wide at base, glabrous except for a few
trichomes near the base. Fruit globose, 3-5
mm high, 4-7 mm wide; seed light brown.
Fig. 2.

Additional selected specimens examined : Queensland.
Wide Bay District: Tinana Creek, 7 km ENE of Tiaro,
Jan 2005, Forster PIF30579 et al. (BRI); NW slopes of
Mt Glastonbury, Dec 1991, Forster PIF9273 & Sharpe
(BRI, CNS, MEL); S of Anderson Road, 10 km W of
Cooroy, Nov 1993, Bean 7065 (BRI, NSW; AD, DNA
n.v.)- Imbil, Dec 1943, Clemens s.n. (BRI [AQ321401]);
near Imbil, Jun 1947, Smith & Webb 31162 (BRI).

264

Moreton District: Scientific Area 2, S.F. 309, Enoggera,
Jan 1992, Forster PIF9353 & Machin* (BRI, CNS,
MEL, NSW; A, B, K, L, NY, MO n.v); Laceys Creek, Mt
Glorious Road, Taylor Range, Feb 1972, Webb & Tracey
11246 (BRI, CANB); The Rafting Ground, Kenmore, Dec
1981, Dickison 294 & Jessup (BRI); Pimpama River, 3.7
km along Ormeau Road from Pacific Highway, c. 11 km
SE of Beenleigh, Jul 1986, Beesley 8497 & Ollerenshaw
(CANB); c. 2 miles [3.2 km] SW of Canungra, Jan 1971,
Williams s.n.* (BRI [AQ310510], CANB, MEL); Mt
Roberts, McPherson Range, Oct 1967, Blake 22848*
(BRI); Roberts Plateau, McPherson Range, Mar 1920,
White s.n.* (BRI [AQ321398]); O’Reilly’s, Lamington
N.P, Nov 1999, Rozefelds s.n.* (HO); Morans Falls,
Lamington N.P, Aug 1978, Floyd s.n.* (CANB289092).
New South Wales. Pacific Highway - Byron Bay Road,
May 1964, Williams s.n. (NSW); Casino, Dec 1908,
Pope s.n. (NSW).

Distribution andhabitat : Pseudoweinmannia
lachnocarpa extends from near Tinana in
south eastern Queensland to Casino in New
South Wales (Map 1). It occurs in riverine,
coastal or montane rainforest from near sea
level to 1000 m.

Phenology : Pseudoweinmannia lachnocarpa
flowers from October through to January;
with fruiting specimens collected between
November through to March.

Typification : Mueller’s (1874) description of
Weinmannia lachnocarpa cited collections
by both C.Moore and W.Guilfoyle from the
Tweed District of northern New South Wales.
The Guilfoyle specimen was initially selected
by Hoogland in 1984 as the lectotype, and
is an appropriate choice as it is the only
fruiting specimen. A specimen in K, which
is also fruiting, was sent by Mueller and it
matches the colouration and appearance of
the lectotype and is interpreted as a putative
isolectotype.

Notes : The type of Pseudoweinmannia
lachnocarpa var. parvifolia F.M. Bailey
differs from the more typical form of the plant
in having smaller leaves and fewer stamens.
More comprehensive collections, particularly
of flowering material, from throughout
the range of the species are needed to fully
assess this variation although there seems
little justification, at present, to recognise a
separate variety.

Etymology : The species name, lachnocarpa ,
comes from the Greek lachnos , wool and

Austrobaileya 8(3): 252-266 (2011)

karpos , fruit, alludes to the dense indumentum
covering the fruits.

Common names applied to this species
include ‘Marara’, ‘Rose Marara’, ‘Mararie’,
‘Merrany’, ‘ScrubRosewood’, ‘RedCarabeen’
(Bailey 1900, 1913; Maiden 1922; Francis
1951; Floyd 1989).

Acknowledgments

This study has benefited from the detailed
notes compiled by the late Ru Hoogland,
although our conclusions regarding the number
of species occurring in Australia differs from
his. We are grateful to an anonymous referee
and Paul Forster for their helpful advice
and comments on an initial draft of this ms.
We thank the Australian Botanical Liaison
Officer, Ken Hill, for arranging cibachromes
of material of both species in Kew. Andrew
Ford and Bernie Hyland, and Robert
Kooyman assisted with fieldwork in 1998 and
1999 respectively. We thank Phillip Brown,
Department of Agriculture, University of
Tasmania for access to a critical point drier
and David Steele, Central Science Laboratory,
University of Tasmania for assistance with
SEM studies.

References

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Forest Flora of New South Wales. 7: 6-8, t. 229.

J. Spence, Government Printer: Sydney.

Matthews, M.L. & Endress, P.K. (2002). Comparative
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morphology of the Australian species of
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Botany 15: 221-236.

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Vol. 4. Inkata Press: Melbourne.

Weberling, F. (1989). Morphology of flowers and
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266

Austrobaileya 8(3): 252-266 (2011)

Map 1. Distribution of Pseudoweinmannialachnocarpa
and P. apetala in eastern Australia.

A taxonomic revision of Nyssanthes R.Br. (Amaranthaceae)

A.R.Bean 1 & C.H.Miller 2

Summary

Bean, A.R. & Miller, C.H. (2011). A taxonomic revision of Nyssanthes R.Br. (Amaranthaceae).
Austrobaileya 8(3): 267-279. The endemic Australian genus Nyssanthes is revised, with the
recognition of four species, N. erecta R.Br., N. diffusa R.Br., N. impervia A.R.Bean sp. nov. and N.
longistyla C.H.Mill. sp. nov. All species are illustrated, and distribution maps provided. Notes for
each species are given on conservation status, distribution and habitat, and an identification key is
presented.

Key Words: Amaranthaceae, Nyssanthes, Nyssanthes impervia , Nyssanthes longistyla , Australia
flora, Queensland flora, taxonomy, identification key, new species

'A.R. Bean, Queensland Herbarium, Department of Environment and Resource Management,
Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia. E-mail: tony.
bean@der m. qld. gov. au

2 C.H. Miller, Australian National Herbarium, Centre for Australian National Biodiversity Research,
G.RO. Box 1600, Canberra 2601, Australian Capital Territory, Australia. E-mail: cathy.miller@csiro.au

Introduction

Nyssanthes R.Br. is a small genus endemic to
eastern Australia. All species are perennial
shrubs or subshrubs characterised by their
opposite leaves, 2-locular anthers, 4-merous
flowers, spreading spinose bracteoles, and
tepals of unequal length. Nyssanthes belongs to
Amaranthaceae Subfamily Amaranthoideae,
but its closest affinities are unclear. It shows
superficial similarities to Achyranthes L.,
but pollen morphology suggests an affinity
with Deeringia R.Br. and Celosia L. (Borsch
1998).

Nyssanthes was named by Brown (1810),
from specimens he collected while in
Australia. Brown described three species,
Nyssanthes diffusa, N. erecta and N. media.
Moquin (1849) maintained all of these, but
Bentham (1870) reduced Nyssanthes media
to synonymy with N. diffusa. Little has been
written about the taxonomy of Nyssanthes
since the work of Bentham. Townsend
(1993) included a description of Nyssanthes
in his comprehensive synopsis of the
Amaranthaceae, although this did not expand
beyond that written by Bentham.

The two common species, Nyssanthes
diffusa and N. erecta , have been treated in
state or regional floras (Ross 1983; Jacobs &

Accepted for publication 28 June 2011

Lapinpuro 1990). They are not infrequent as
a component of the understorey in woodlands
and forests of eastern Queensland and New
South Wales. Despite being indigenous,
they are often termed “weeds”, because
they are prickly and may proliferate under
optimal conditions. Two additional species
have recently been recognised from western
Queensland, and they are newly described
here as Nyssanthes impervia A.R.Bean and
N. longistyla C.H.Mill. They inhabit shallow
or skeletal soils in areas that receive low
annual rainfall (280-445 mm).

Discussion of some morphological
characters

Due to the lack of basic knowledge of
Nyssanthes morphology, a number of
misconceptions have been propagated in the
literature. Townsend (1993) mentioned that
all flowers are hermaphroditic in Nyssanthes.
However, the basal flowers formed in the leaf
axils of Nyssanthes diffusa are sterile. They
usually comprise a bract, two bracteoles,
and one tepal. All of the other tepals are
rudimentary or absent, and no male or female
parts can be observed. These sterile flowers
seem to develop in all leaf axils and persist
year-round. Subsequently (and perhaps at
certain times of the year) fully fertile and
bisexual flowers develop. As far as is known,
sterile flowers do not occur in the other
species.

268

Austrobaileya 8(3): 267-279 (2011)

The length of the bract, bracteoles and the
longest tepal vary greatly within each species
of this genus, as Bentham (1870) observed.
However, the variation is not random. The
sterile flowers of Nyssanthes diffusa have
very long spinose bracts and bracteoles, and
sometimes the longest tepal is also of similar
length and is spinose. Subsequent flowers in
an inflorescence display a gradual decrease
in the lengths of the bracts and bracteoles,
even though the change from sterile to fertile
flowers is apparently rapid and without
intermediate forms.

Jacobs & Lapinpuro (1990) stated that
stamen number varies within species of this
genus. We have not found this to be the case.
All flowers of Nyssanthes erecta examined,
have possessed four fertile stamens, and for
N. diffusa consistently two. However, the
anthers are very easily shed, and an erroneous
stamen count can easily occur if the flower is
not opened very carefully.

Materials and methods

This revision is based upon field examination
of all species, and herbarium holdings at AD,
BRI, CANB, CNS, HO, MEL, NSW and
NT, and type specimens from BM and K.
Measurements of leaves, bracts and bracteoles
were made from dried herbarium specimens;
measurements of tepals, stamens and styles
were made from material preserved in spirit
or reconstituted with boiling water.

Common abbreviations in the specimen
citations are N.R (National Park) and S.F.
(State Forest).

Taxonomy

Nyssanthes R.Br., Prodr. 418 (1810).

Type species: N. diffusa R.Br. (lecto: here
designated).

Perennial herbs or shrubs. Leaves opposite,
entire, exstipulate. Inflorescence spicate, in
leaf axils. Rachis present or absent. Flowers
sessile. Bract 1 per flower, bracteoles 2 per
flower, spinose; basal parts cymbiform, more
or less appressed. Sterile flowers sometimes
present. Fertile flowers all bisexual, sessile.
Perianth 4-merous, tepals cymbiform,
heteromorphic, the outermost (posterior) tepal
being the longest, the next (anterior) tepal
being shorter; the two inner tepals shorter
again, but about the same size and shape as
each other. Stamens 2 or 4; anthers bilocular.
Pseudostaminodes present, 4. Ovary 1-locular,
1-ovular. Style not lobed, capitate. Fruit an
indehiscent utricle, dispersed as a unit with
perianth and bracteoles. Seeds 1, ellipsoid to
ovoid.

Etymology : From the Greek nyssa (a prickle),
and anthos (flower), in reference to the prickly
bracteoles and bracts found in all species of
the genus.

Typification: In Brown’s diary the description
of Nyssanthes diffusa has been annotated with
“nov: gen Amaranthor: Infs:”. Additionally the
name was written as “Nyssanthos axillaris”
but this has been crossed out and replaced by
“Nyssanthes diffusa” while the other species
names had not been altered. This would
indicate that Brown had determined the name
of this taxon before he named the other two
taxa (J.G.West pers. comm). For these reasons
Nyssanthes diffusa R.Br. has been selected as
the lectotype species for the genus.

A genus of four species endemic to eastern
Australia.

Bean & Miller, Revision of Nyssanthes

269

Key to the species of Nyssanthes

1 Leaf lamina 2.5-5.5 times longer than broad; leaf apical mucro absent or

up to 0.7 mm long; longest tepal recurved; central Queensland, eastern

New South Wales.1. N. erecta

1. Leaf lamina 1.5-2.5 times longer than broad; leaf apical mucro 0.5-2.1

mm long; longest tepal straight.2

2 Fully expanded leaves sparsely tomentose; inflorescence up to 22-flowered;

stamens two; eastern New South Wales & Queensland.2. N. diffusa

2. Fully expanded leaves moderately to densely tomentose; inflorescence

2-5(-8) flowered; stamens four.3

3 Style 0.6-0.8 mm long; pseudostaminodes without dorsal appendage;

bracts 0.8-1.2 mm wide; southwest Queensland.3. N. impervia

3. Style 27-3.5 mm long; pseudostaminodes with trifid dorsal appendage;

bracts 1.4-2 mm wide; southwest Queensland.4. N. longistyla

1. Nyssanthes erecta R.Br., Prodr. 418
(1810). Type citation: “(J.) v.v.” Type: New
South Wales. Banks of the Nepean [River] on
Badgery’s farm, [December 1804 or January
1805], R. Brown [Bennett No. 3070] (lecto:
BM000795238, here designated; isolecto:
BM000795214).

Dichasially branched perennial forb or
shrub to 0.8 m high. Stems striate, grooved,
sparsely tomentose; hairs antrorse, appressed
to spreading, 0.1-0.5 mm long. Petioles
1-5 mm long; lamina narrowly-elliptic
to narrowly ovate, 14-26 mm long, 3.5-8
mm wide, 2.5-5.5 times longer than broad,
sparsely tomentose when fully expanded;
hairs unicellular, appressed to spreading,
0.15-0.5 mm long; base attenuate, apex
acute or mucronate (mucro 0-0.7 mm long).
Inflorescence up to 16-flowered. Rachis to
8 mm long, sparsely to densely tomentose.
All flowers fertile. Bract 1.3-1.75 mm long,
1-1.3 mm wide; apex acuminate, spinulose,
acumen 0.2-1.5 mm long; base broadly ovate,
hyaline; remaining firmly attached to rachis
upon disarticulation of flower. Bracteoles
2.5-4.5 mm long, 0.9-1.7 mm wide; basal
part ovate to broadly ovate, moderately
tomentose, with thick, yellowish-brown
midrib, otherwise hyaline; acumen slender,
patent, yellowish-brown, spinose; remaining
attached to perianth upon disarticulation of
flower. Perianth 1.4-2 mm wide. Longest
tepal lanceolate-acuminate, 5.5-77 mm
long, recurved, basal half very sparsely to

densely tomentose; midrib prominent, with
1-2 pairs of lateral longitudinal ribs visible;
apex spinose. Next longest (anterior) tepal
slightly shorter but otherwise similar, 4.2-
6.8 mm long. Inner 2 tepals narrowly-ovate,
2.8-3.8 mm long, glabrous; apex acuminate,
spinulose. Fertile stamens 4, filaments
slender, 0.4-07 mm long; anthers 0.35-0.5
mm long, white to yellow. Staminodes absent.
Pseudostaminodes quadrate, glabrous, almost
filling space between filaments, 0.3-0.8
mm high, translucent; distal margin entire
or ciliate; dorsal appendage absent. Ovary
oblate, c. 0.8 mm long, glabrous. Style simple,
0.4-0.6 mm long, stigma capitate. Utricle
with membranous covering, irregularly
fragmenting. Seed obloid, 0.75-1.6 mm long,
0.65-1.1 mm wide, brown. Fig. 1.

Additional selected specimens examined: Queensland.

South Kennedy District: ‘Logan Downs’, Clermont,
Jun 1942, Fairbairn s.n. (BRI [AQ683973]). Mitchell
District: Jericho, Apr 1946, Clemens s.n. (BRI
[AQ683970]); ‘Gartmore’ on Alpha - Tambo road, c.
11 km NNE of Tambo, Apr 1961, Johnson 2185 (BRI).
Leichhardt District: ‘Minerva’, Mar 1935, Blake
7946 (BRI, CANB); ‘Cluden’, south of Taroom, Oct
1954, Johnson & Pedley 17 (BRI); ‘Nirvana’, c. 15
km WNW of Banana, Apr 2003, Bean 20152 (BRI);
SE of ‘Redcliffe’ via Baralaba, Mar 2005, Bean 23609
(BRI). Port Curtis District: Charons Ferry Station
near Styx, Jun 1942, Blake 14606 (BRI); Rosedale,
N.C. [North Coast] Line, Nov 1932, Dovey 164 (BRI);
Experimental Farm, Biloela, Oct 1947, Smith 3550
(BRI). Burnett District: on property of H.J. Brunje,
West Goomeri, Mar 1962, Saint-Smith s.n. (BRI
[AQ178268]). Warrego District: 51 kmN of Charleville
along road to Augathella, W side of road. Mar 1976,
Purdie & Boyland 163 (BRI). Darling Downs District:

270 Austrobaileya 8(3): 267-279 (2011)

Fig. 1. Nyssanthes erecta. A. upper part of flowering plant x l.B. leafy branchlet x 1.5. C. complete flower x 8. D. bract
x 12. E. bracteole x 12. F. bracteole x 12. G. outermost tepal x 12. H. second tepal x 12.1. third tepal x 12. J. innermost
tepal x 12. K. ovary, style, stamens and pseudostaminodes x 16. L. seed x 16. A, C from Bean 20152 (BRI); B from
Bean 23609 (BRI); L from Clemens s.n. (BRI [AQ683970], Del. W.Smith.

Bean & Miller, Revision of Nyssanthes

Chinchilla, Portion 100, ‘Mylo’, Chinchilla Shire, Dec
1989, Mann s.n. (BRI [AQ467453]); Kindon Station,
about 54 miles [90 km] NNE of Goondiwindi, Dec 1938,
Smith 610 (BRI); ‘Woodside’, Pratten, Oct 1962, Walsh
s.n. (BRI [AQ169899]); Bybera, Jan 1934, White 9700
(BRI); Wallumbilla, May 1916, White 9984 (BRI); c. 0.5
km S of Pelican Hall on Chinchilla - Burra Burri Road,
Apr 1981, Wilson 3395 (BRI, NSW). Moreton District:
Crows Nest, Mar 1948, Clydesdale 87 (BRI). New South
Wales. ‘Roma’, near Moree, May 1977, Bradbrook
s.n. (NSW); Stoney Creek Road at South Creek, south
of ‘Berkshire Park’, Dec 1982, Coveny et al. 11473
(NSW); Tenterfield, s.dat., s.coll. (MEL 59864); Bank
of Shoalhaven River, Nowra, Feb 1933, Menaghan s.n.
(NSW); Bullen Buttress, Shoalhaven Gorge, c. 8 km
upstream from the Tallowa Dam site, Feb 1974, Pullen
8775 (AD, CANB, HO, NSW); New England, s.dat.,
Stuart s.n. (MEL59863).

Distribution and habitat : Nyssanthes erecta
is widespread from the Mt Coolon area (west
of Mackay) in Queensland south to Nowra in
New South Wales (Map 1). It grows on heavy
clay soils, often in association with brigalow
(Acacia harpophylla F.Muell. ex Benth.) or
poplar box (Eucalyptus populnea F.Muell.),
but also in a range of other eucalypt-dominated
communities.

Phenology: Flowers and fruits may be found
throughout the year.

Typification: Brown’s diary does not specify
any locality for Nyssanthes erecta. However,
one of the sheets at BM (000795238) has
the locality written in Brown’s handwriting.
Also written next to the name is “Prod. 418”
indicating that Brown probably saw this
specimen. This sheet has been designated
as the lectotype. There is another sheet at
BM (000795214) that matches the lectotype
material but does not have a label in Brown’s
handwriting and with a general location of
Port Jackson. This was probably from the
same collection but has been separated as
there is no Bennett number (J.G. West pers.
comm). This specimen is here designated as
an isolectotype.

Notes: The conspicuously recurved perianth
exhibited by this species is not found in any
other species of Nyssanthes. In all of the other
species, the perianth is straight or slightly
incurved.

Conservation status: A widespread and
relatively common species.

271

2. Nyssanthes diffusa R.Br., Prodr. 418 (1810).
Type: [Queensland. Port Curtis District:].
Pine Mountain, Shoalwater Bay, 3 September
1802, R.Br own [Bennett No. 3069] (holo: BM;
iso: E n.v., K000357277 & 000357276).

N. media R.Br., Prodr. 418 (1810). Type
citation: “(J.) v.v” Type: New South Wales.
Port Jackson, banks of the Hawkesbury,
R.Brown s.n. (lecto: BM000795233, specimen
on right hand side of sheet, here designated;
isolecto: BM000795234 & 000795235).

Divaricately branched herb or shrub to 0.7
m high. Stems striate, grooved, sparsely
tomentose, glabrescent; hairs antrorse,
appressed to spreading, 0.2-0.4 mm long.
Petioles 0.5-3 mm long; lamina broadly
elliptical, 8-35 mm long, 4.5-18 mm wide,
1.5-2.5 times longer than broad, sparsely
tomentose when fully expanded; hairs
unicellular, appressed, 0.2-0.5 mm long; base
cuneate, apex mucronate (mucro 0.5-1.7 mm
long). Inflorescence up to 22-flowered. Rachis
to 10 mm long, with simple non-ciliolate
hairs. Up to 4 basal flowers sterile, subsequent
flowers fertile. Sterile flowers: Bract subulate
throughout and spinose, remaining attached
to perianth upon disarticulation of the
flower, 3.5-7 mm long; bracteoles subulate
throughout and spinose, 5-9 mm long;
longest tepal somewhat expanded at the base,
3-7mm long; all other tepals rudimentary
or absent; stamens and ovary absent. Fertile
flowers: Bract subulate, 1.3-4.5 mm long,

O. 5-1.2 mm wide; basal part ovate to broadly
ovate, hyaline, 0.6-1.5 mm long; acumen 1-
3.5 mm long, spreading, spinose; remaining
attached to rachis upon disarticulation of
flower. Bracteoles 2.1-4.6 mm long, 0.6-1.2
mm wide; basal part ovate to broadly ovate,
with thick, yellow-brown midrib, otherwise
hyaline, sparsely tomentose; acumen slender,
patent, 1-5 mm long, yellowish-brown,
spinose; remaining attached to perianth upon
disarticulation of flower. Perianth 0.9-1.4 mm
wide. Longest (posterior) tepal lanceolate-
acuminate, 5-8.7 mm long, straight, basal half
tomentose on outer surface; midrib prominent,
with 2 or 3 pairs of lateral longitudinal ribs;
apex subulate, spinose. Next longest (anterior)
tepal 2.6-4.9 mm long, straight, sparsely
tomentose on outer surface; apex acute,

272

spinulose. Inner 2 tepals narrowly-ovate, 2.1-
2.9 mm long, sparsely tomentose or glabrous
on outer surface; apex acute, not spinose.
Fertile stamens 2, filaments slender, 0.7-1.1
mm long, often differing in length in the same
flower; anthers 0.25-0.4 mm long, white to
yellow. Staminodes 0-2, 0.5-0.8 mm long,
shorter and thinner than stamen filaments.
Pseudostaminodes quadrate, translucent,
0.4-0.8 mm high, almost filling the space
between filaments, glabrous, distal margin
entire or denticulate, dorsal appendage absent.
Ovary lenticular, 0.3-0.6 mm long, glabrous.
Style simple, 0.3-0.5 mm long, stigma
capitate. Utricle with membranous covering,
irregularly fragmenting. Seed ovoid, 1-1.5
mm long, 0.9-1.1 mm wide, 0.7-0.8 mm deep,
brown. Fig. 2.

Additional selected specimens examined : Queensland.
North Kennedy District: 3 km along Tumoulin Road
from Kennedy Highway Junction, Jan 2004, McDonald
1692 (BRI). South Kennedy District: Eungella Range,
Sep 1938, White 12979 (BRI). Leichhardt District:
Crows Apple Scrub, ‘Rookwood’, Apr 1991, Forster
P1F7977&McDonald(BRiy, Rosedale near Baralaba, Sep
1959, Johnson 920 (BRI). Brigalow Research Station, 20
miles [33 km] NW of Theodore, Mar 1963, Johnson 2632
(BRI); Auburn Range S.F., about 50 km S of Thangool,
Jun 1996, Bean 10363 (BRI); road to Sandstone Bore,
S.F. 46, NW of Taroom, Jan 2003, Bean 19921 (BRI).
Port Curtis District: Bulburin S.F. 67, vicinity of Forest
station, Apr 1980, McDonald et al. 3228 (BRI, CANB).
Burnett District: Bunya Mountains. Near Mowbullan
guest house, Jun 1961, Willis s.n. (MEL). Wide Bay
District: Stony Creek, 4 km E of Didcot, Biggenden
Shire, Dec 1984, Forster P1F1969 (BRI); Glastonbury
district; c. 24 km WSW of Gympie, Apr 1978, Henderson
2623 (BRI). Darling Downs District: Teviot Range, 35
1cm NE of Killarney (SW of Wilsons Peak), May 1971,
Briggs 4210 (NSW). Moreton District: 10 km E of Mt
Nebo, Nov 1980, Dillewaard& Olsen 241 (BRI, CANB);
Table Top, Main Range, Toowoomba, May 1937, Roe
A27 (CANB); Lamington N.P, near turnoff to Moran’s
Falls from O’Reilly’s guest house. May 1981, Willis s.n.
(MEL). New South Wales. 0.5 km along Tucker Box
Road, Breury S.F., 20 km NNW of Urbenville, Nov
1987, Coveny et al. 12810 (CANB, NSW); ‘The Native
Vineyard’ (near Cobbity Trig), 5.5 km N of Cobbity,
Mar 1976, Coveny et al. 7445 (NSW); Waa Gorge,
Mt Kaputar N.P, 68 km NE of Narrabri by road, Nov
1976, Coveny & Roy 9019 (NSW); Yellow-Rock Road,
about 4 miles [6.6 km] SW of ‘Albion Park’, Feb 1967,
Evans 2584 (BRI, CANB, MEL, NSW); Berry, Jan 1929,
George s.n. (NSW); Acacia Creek, Jan 1956, Gray 3787
(CANB); ‘Taroona’, south of Bowling Alley Point, Mar
1995, Hosking 1096 (CANB); Scone, May 1902, Maiden
& Boorman s.n. (NSW); ‘Bajoloma’, Cowal Creek Road,
Bellangry, Sep 1987, Miller & Whaite 458 (CANB);
Between Kiama and Jamberoo, South Coast, Mar 1964,

Austrobaileya 8 ( 3 ): 267-279 ( 2011 )

Pullen 4028 (CANB, NSW); c. 8 km SW of Beelbrook
on Mackenzies Creek Road, Apr 1976, Wilson 1564
(NSW); 5.8 km W of Bingara on Narrabri Road, May
1985, Wilson 6232 (BRI, NSW).

Distribution and habitat : Nyssanthes diffusa
extends from the southern coast of New South
Wales to Ravenshoe in north Queensland.
While most occurrences are within 200 km
of the coast, it does extend as far inland as
Carnarvon Station, south-west of Springsure
(Map 2). It inhabits sunny spots (e.g. along
tracks) in rainforest (especially the Araucarian
vine forests), and sometimes in eucalypt
open forest. When it occurs in eucalypt
communities, it is usually found in a shady
microhabitat.

Phenology : Flowers and fruits may be found
at any time of the year.

Typification : There are three sheets of
Nyssanthes media located at BM. The sheet
BM000795233 has a reference to Brown’s
Prodromus written on the label in Brown’s
handwriting attached to the specimen on
the right hand side of the sheet and is here
designated as the lectotype. The specimen
on the left hand side of the sheet has a
different location (Mount Hunter) and is a
lusher specimen than normal and has been
designated as a residual syntype. The other
two BM sheets (000795234 and 000795235)
do not have labels in Brown’s handwriting
but have other labels indicating Port Jackson
and match the lectotype material (J.G. West
pers. comm.). These are here designated as
isolectotypes.

Notes : The basal (first-formed) flowers of the
inflorescence in this species are usually sterile.
The bract and bracteoles and the posterior
tepal are all long and rigid, and the latter
has only a small hyaline swelling at its very
base. The other tepals are much reduced, and
there are no male or female parts. Bentham
(1870) wrote “sometimes all three bracts are
subulate almost from the base and 3 to 4 lines
long”. This is a reference to the sterile flowers,
although he did not record them as such.

Nyssanthes media has been maintained in
synonymy here following Bentham (1870) as
there are not enough characters to distinguish
it from N. diffusa. Brown (1810) separated

Bean & Miller, Revision of Nyssanthes

273

Fig. 2. Nyssanthes diffusa. A. branchlet with sterile flowers x 2. B. sterile flower x 6. C. branchlet with fertile flowers
x 1.5. D. inflorescence with fertile flowers x 3. E. fertile flower x 12. F. bract x 12. G. bracteole x 12. H. outermost tepal
x 12.1. second tepal x 12. J. third tepal x 12. K. innermost tepal x 12. L. style, stamens and pseudostaminodes x 24. M.
seed x 24. A, B from Bean 19921 (BRI); C-L from Bean 10363 (BRI); M from Johnson 920 (BRI). Del. W.Smith.

274

the two taxa by the number of nerves on the
tepals, the level of pubescence of the plant
and the awn to leaflet ratio. Examination of
the type specimens found that the first two
characters varied considerably and could not
be used to consistently separate the taxa. The
third character is not useful as it is not clear
from his descriptions as to his definition of an
awn and several interpretations are possible.
Bentham (1870) stated that ‘W. media appears
to me to be rather a luxuriant state than a
variety of N. diffusa ”.

Conservation status: A widespread and
relatively common species.

3. Nyssanthes impervia A.R.Bean species
nova afinis N. diffusae , sed foliis hirsutioribus,
inflorescentiis 3-floris, et staminibus fertilibus
4 antheris 0.2-0.3 mm longis, distinguenda.
Typus: Queensland. Gregory South District:
4 km (direct) NNW of‘Trinidad’ Homestead,
E ofWindorah, 1 September 2010, A.R. Bean
30219 (holo: BRI; iso: CANB, K, MEL,
NSW).

Divaricately branched, woody shrub up
to 1 m high. Stems obscurely grooved or
terete, young growth moderately to densely
tomentose; hairs antrorse, appressed to
spreading, 0.15-0.4 mm long. Petioles 0-2
mm long. Lamina broadly elliptical, 5-11 mm
long, 2.5-6 mm wide, 1.8—2.2 times longer
than broad, with dense hairs on both surfaces
when fully expanded; hairs unicellular,
appressed, 0.25-0.6 mm long; base cuneate,
apex mucronate (mucro 0.7-2.1 mm long).
Inflorescences 3-flowered; rachis absent.
All flowers fertile, lateral flowers maturing
before central flower. Bract 6-8.5 mm long,
0.8-1.2 mm wide, remaining attached to
stem upon disarticulation of fruit; basal part
ovate, hyaline, 1.2-1.6 mm long, sparsely
tomentose; upper part subulate, 4.5-7 mm
long. Bracteoles 5.2-7.8 mm long, 1.5-2 mm
wide; basal part ovate, hyaline, 1.3-1.8 mm
long, acumen 4.2-6.5 mm long, spreading,
spinose. Perianth 1.2-1.8 mm wide. Longest
tepal lanceolate-acuminate, 6.5-9.5 mm long,
straight, glabrous, midrib prominent and 2 or 3
pairs of lateral ribs visible; apex spinose. Next
longest tepal lanceolate-acuminate, 5-6.8
mm long, with dense, appressed to spreading

Austrobaileya 8(3): 267-279 (2011)

tomentum on outer surface, apex acute, not
spinose. Inner 2 tepals deflate, 3.5-6.2 mm
long, with moderately dense, appressed to
spreading tomentum on outer surface, apex
acute, not spinose. Fertile stamens 4, filaments
0.2-0.45 mm long; anthers 0.2-0.3 mm long,
white. Staminodes absent. Pseudostaminodes
4, quadrate, c. 0.3 mm long, distal margin
irregular, dorsal appendage absent. Ovary
oblate, c. 0.25 mm long, glabrous. Style
simple, 0.6-0.8 mm long, stigma capitate.
Fruits not seen. Fig. 3.

Additional specimens examined: Queensland. Gregory
South District: 4 km (direct) NNW of ‘Trinidad’
Homestead, E of Windorah, Sep 2010, Bean 30228
(AD, BRI, US); Cheviot Range, c. two-thirds of the
way between ‘Budgerygar’ Homestead and ‘Trinidad’
Homestead (towards Trinidad), Aug 1997, Purdie 4598
(BRI, CANB).

Distribution and habitat: Nyssanthes
impervia is currently known only from the
type location in south-western Queensland
(Map 3). It inhabits shallow sandy soils at the
base of rock outcrops or cliffs, in low open
woodland dominated by Acacia catenulata
C.T.White, A. shirleyi Maiden or Eucalyptus
thozetiana F.Muell. ex R.T.Baker. Other
associated shrubs include Eremophila spp.,
Ptilotus obovatus and Sida spp.

Phenology: Flowers are recorded in August
and September; fruits have not been
collected.

Notes: Nyssanthes impervia is similar to N.
diffusa , but differs by the denser indumentum
of the leaves, the 3-flowered inflorescences
(up to 22-flowered for N. diffusa ), the 4 fertile
stamens in each flower (2 fertile stamens
for N. diffusa), and the anthers 0.2-0.3
mm long (0.25-0.4 mm for N. diffusa). It is
similar stature and appearance to Nyssanthes
longistyla. However, Nyssanthes impervia
differs by the much shorter style (0.6-0.8 mm
long), and the pseudostaminodes that lack a
dorsal appendage.

Conservation status: There are 300-400
plants of this species currently known at the
type locality. Feral goats are present in the
area (J. Silcock pers. comm. Nov 2010) and
the effects of their grazing were evident on
most plants in September 2010. Under the

Bean & Miller, Revision of Nyssanthes

275

Fig. 3. Nyssanthes impervia. A. flowering branchlet x 3. B. immature flower x 9. C. bract x 12. D. bracteole x 12.
E. bracteole x 12. F. outermost tepal x 12. G. second tepal x 12. H. style, stamens, pseudostaminodes and innermost
tepal x 24.1. third tepal x 12. J. innermost tepal x 12. A-E from Purdie 4598 (BRI); F-J from Bean 30219 (BRI). Del.
W. Smith.

276

criteria of the IUCN Red List (IUCN 2001), a
conservation status of Vulnerable is suggested
[VU Blab(v) + 2ab(v); Cl+2a(i,ii); Dl+2],

Etymology : From the Latin impervius
meaning impassable. This is in reference to
the very spiny nature of the plant, making it
impervious to human touch and impassable to
human traffic.

4. Nyssanthes longistyla C.H.Miller species
nova distinguenda a N. erecta habitu lignea;
tepalis duobus exterioribus pilis extus dense
obtectis; tepalis interioribus, filamentis
staminalibus, antheris et stylo longioribus;
pseudostaminodiis multo longioribus,
laciniatis et fimbriatis, dorsaliter squama
trifida praeditis; fructibus multo latioribus.
Typus: Queensland. Gregory North District:
Mt Booka Booka, Elderslie Station, c. 6 km E
of Diamantina River, Boulia - Winton road,
10 June 1997, L.A.Craven, C.L.Brubaker &
J.P. Grace 9915 (holo: CANB; iso: AD, BRI,
DNA, K, MEL, MO, NSW).

Dichasially branched woody shrub up to 1.2
m tall. Stems striate, grooved, on older stems,
young growth densely tomentose; hairs
antrorse, appressed to spreading, 0.3-0.6
mm long. Petioles 0.5-3.5 mm long. Lamina
elliptical, 6-15 mm long, 3-8.5 mm wide,
2-2.5 times longer than broad, moderately
tomentose on both surfaces when fully
expanded; hairs unicellular, appressed to
spreading, 0.5-1 mm long; base attenuate to
cuneate, apex mucronate, (mucro 0.5-1.3 mm
long). Inflorescence with (2-) 5-8 flowers;
rachis to 1 mm long. All flowers fertile. Bract
3.2-6.5 mm long, 1.4-2 mm wide, remaining
firmly attached to rachis upon disarticulation
of fruit; basal part ovate to broadly ovate,
hyaline, 1.2-2.5 mm long, becoming recurved
with maturity, glabrous or with a few hairs on
back near base; upper part subulate, 2.5-5.8
mm long. Bracteoles 4-7.2 mm long, 1.7—
2.3 mm wide; base ovate to broadly ovate,
hyaline, 1.5-2.8 mm long, acumen 2.5-6.5
mm long, spinose; moderately tomentose
along edge of nerve, remaining attached
to perianth upon disarticulation of flower.
Perianth 1.5-2.2 mm wide. Longest (posterior)
tepal lanceolate-acuminate, 6.5-9 mm long,
straight, sparsely tomentose along edge of

Austrobaileya 8(3): 267-279 (2011)

nerve of top half; midrib prominent and 3-5
pairs of lateral longitudinal ribs visible; apex
spinose. Next longest (anterior) tepal similar,
5.5-7.5 mm long, densely tomentose. Inner
2 tepals narrowly-ovate, 5-6.5 mm long,
glabrous; apex acuminate. Fertile stamens 4,
filaments slender, 2.6-3.3 mm long; anthers
1.2-1.7 mm long, cream. Staminodes absent.
Pseudostaminodes 4, quadrate, 1.5-2 mmhigh,
distal margin laciniate, fimbriate translucent,
dorsal appendage trifid with ciliate margins,
0.6-1.1 mm long. Ovary obconical, c. 0.8 mm
long, glabrous. Style simple, 27-3.5 mm long,
stigma capitate. Utricle with membranous
covering, irregularly fragmenting. Seed
ellipsoid, 1.8-2.1 mm long, 1.4-1.5 mm wide,
brown. Fig. 4.

Additional specimens examined : Queensland:

Mitchell District: Mt Felix, c. 90 km N of Jundah on
road to Winton, May 2010, Bean 29779 & Emmott (BRI,
CANB, MEL, MO). Gregory North District: On upper
slopes of Mt Booka Booka, ‘Elderslie’, Oct 1935, Blake
10056 (BRI, NT); Mt Booka Booka, ‘Elderslie’, c. 70 km
W of Winton, Aug 2009, Silcocks.n. (BRI [AQ785536]).

Distribution and habitat : Endemic to
Queensland, where it is known from two
locations in the west of the state (Map 3).
It grows on the upper slopes of mountains
in gravelly shaly soil in association with
other shrubby genera including Senna and
Abutilon.

Phenology : Flowers and fruits have been
recorded in May, August and October.

Notes : This species varies considerably from
the other species of Nyssanthes. It differs from
Nyssanthes erecta by being a woody shrub
sometimes more than a metre tall, densely
covered in hairs all over; leaves are small
and elliptic; the tepals are usually densely
covered with hairs on the outside; the fruits
are much broader; style, staminal filaments,
anthers and pseudostaminodes are much
longer. It is the only Nyssanthes species to
have a trifid appendage on the dorsal surface
of the pseudostaminode. It and Nyssanthes
impervia have a scattered distribution in
inland Queensland, compared to the more
extensive distribution of N. erecta and
N. diffusa throughout large areas of eastern
Queensland and New South Wales.

Bean & Miller, Revision of Nyssanthes

277

Fig. 4. Nyssanthes longistyla. A. flowering branchlet x 4. B. flower at anthesis x 8. C. bracteole x 12. D. bract x 12.
E. outermost tepal x 12. F. second tepal x 12. G. third tepal x 12. H. innermost tepal x 12.1. ovary, style, stamens and
pseudostaminodes x 12. J. pseudostaminode x 16. A-C, E-J from Bean 29779 & Emmott ( BRI); D from Silcock s.n.
(BRI [AQ785536]). Del. W.Smith.

278

Conservation status : There are between
100 and 200 plants present of this species
on Mt Booka Booka (J.Silcock pers. comm.
Aug 2009), and all plants are very heavily
grazed by feral goats. At Mt Felix, there are
more than 1000 plants; here all plants are
in good condition and feral goats are not
present. Applying the criteria of the IUCN
Red List (IUCN 2001), a conservation status
of Vulnerable is proposed. [VU Blab(v) +
2ab(v); Cl; D2],

Etymology : From the Latin longus meaning
long and stylus meaning style in reference to
its long style. The other Nyssanthes taxa have
styles that are 0.3-0.8 mm long, while in N.
longistyla the style is 27-3.3 mm long.

Acknowledgements

We are grateful to Will Smith (BRI) for the
excellent illustrations, Laurie Adams (CANB)
and Peter Bostock (BRI) for the Latin
diagnoses and Nunzio Knerr (CANB) for the
maps. The first author thanks Jenny Silcock
for surveying the Nyssanthes population
on Mt Booka Booka, and Angus Emmott
for assisting with the visit to Mt Felix. The
second author thanks Lyn Craven (CANB) for
visiting Mt Booka Booka and bringing back
specimens to study and Judy West (CANB)
for assistance with nomenclatural matters.

References

Bentham, G. (1870). Nyssanthes. In Flora Australiensis
5: 246-248. L. Reeve & Co.: London.

Borsch, T. (1998). Pollen types in the Amaranthaceae:
morphology and evolutionary significance.
Grana 37: 129-142.

Brown, R. (1810) [1963 fascimile], Nyssanthes. In
Prodromus florae Novae Hollandiae et Insulae
van-Diemen, p. 418. J.Cramer: New York.

Iucn (2001). IUCN Red List Categories: version 3.1.
IUCN Species Survival Commission. IUCN:
Gland, Switzerland.

Jacobs, S.W.L. & Lapinpuro, L. (1990). Amaranthaceae.
In G. Harden (ed.), Flora of New South Wales 1:
248-60. University of New South Wales Press:
Sydney.

Moquin, C.H. (1849). Amaranthaceae. In A.P de
Candolle (ed.), Prodromus Systematis Naturalis
Regni Vegetabilis 13(2): 231-424.

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Ross, E.M. (1983). Amaranthaceae. In T.D. Stanley
& E.M. Ross (eds.). Flora of south-eastern
Queensland 1: 135. Department of Primary
Industries: Brisbane.

Townsend, C.C. (1993). Amaranthaceae. In K. Kubitzki
(ed ). Families and Genera of Vascular Plants 2:
70-91. Springer-Verlag: Berlin.

'a

Map 1. Distribution of Nyssanthes erect a

Bean & Miller, Revision of Nyssanthes

279

13

Map 2. Distribution of Nyssanthes diffusa

Map 3. Distribution of Nyssanthes impervia m and
N. longistyla ▲

A taxonomic revision of Pterocaulon section Monenteles
(Labill.) Kuntze (Asteraceae: Inuleae-Plucheinae)

A.R. Bean

Summary

Bean, A.R. (2011). A taxonomic revision of Pterocaulon section Monenteles (Labill.) Kuntze
(Asteraceae: Inuleae-Plucheinae). Austrobaileya 8(3): 280-334. Pterocaulon section Monenteles,
comprising all species of the genus from Asia, Malesia, Australia and Melanesia, is revised. 14
species are recognised, including the six new species, P. brachyanthum A.R.Bean, P. ciliosum
A.R.Bean, P. discolor A.R.Bean, P. paradoxum A.R.Bean, P. tricholobum A.R.Bean and P. xenicum
A.R.Bean, and one new combination, P. intermedium (DC.) A.R.Bean. Descriptions, illustrations,
and distribution maps are provided for all taxa. Monenteles sphacelatus and M. spicatus are
lectotypified. The ecology, chemistry and diagnostic morphological characters of members of the
section are discussed, and an identification key to species is provided.

Key Words: Asteraceae, Pterocaulon, Pterocaulon brachyanthum, Pterocaulon ciliosum, Pterocaulon
discolor , Pterocaulon intermedium, Pterocaulon paradoxum, Pterocaulon tricholobum, Pterocaulon
xenicum, taxonomy, Australia flora. New Caledonia flora, Malesia flora, coumarins, identification
key, taxonomy, new species

A.R.Bean, Queensland Herbarium, Department of Environment and Resource Management,
Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: tony.
bean@der m. qld. gov. au

Introduction

Pterocaulon Elliott comprises 26 species
distributed in North America, South America,
south-east Asia, Malesia, Melanesia and
Australia. In a revision of the genus by Cabrera
& Ragonese (1978), four taxonomic sections
were recognised. Three of these sections
consist wholly of American species while
the fourth, Pterocaulon section Monenteles
(Labill.) Kuntze, comprises all of the Old
World taxa.

Monenteles Labill. was described at
generic rank by Labillardiere (1825), with two
species, M. sphacelatus Labi 11. andM spicatus
Labill, both named from specimens collected
in New Caledonia. De Candolle (1836) and
Bentham (1867) maintained Monenteles at
generic rank, but the latter commented, “From
the American genus Pterocaulon it scarcely
differs in the hermaphrodite flowers usually
reduced to a single one”.

The reduction of Monenteles to synonymy
with Pterocaulon was effected by Bentham
(1873), and species combinations were made
by Mueller (1882). Monenteles was reinstated
as a section of Pterocaulon by Kuntze (1903).

Anderberg (1989) suggested that the genus
Monenteles could be resurrected, but later
(Anderberg 1991) he abandoned the idea.

The differences between Pterocaulon
section Pterocaulon and P. section Monenteles
are minimal. Nesom (2006) stated that species
in Pterocaulon section Pterocaulon have 1-17
functionally staminate florets per capitulum,
a 1 or 2-seriate pappus, and the hairs have a
relatively long aseptate portion arising from
a cluster of basal cells; while species in P.
section Monenteles have a single staminate
floret per capitulum, a uniseriate pappus, and
the hairs are more-or-less uniformly septate
throughout.

Pterocaulon has been consistently placed
in the Pluchea-growp of genera. The rank of
this group was originally a Subtribe (Subtribe
Plucheinae Benth. & Hook.f. of the Tribe
Inuleae Cass.; Bentham 1873); then it was
deemed a Tribe (Tribe Plucheeae A.Anderb.;
Anderberg 1991, 1994). Evidence from

molecular data now favours subtribal rank
(Anderberg et al. 2005). Within the subtribe,
Anderberg (1991) grouped Pterocaulon
alongside Neojeffreya Cabrera and
Stenachaenium Benth. in the “Pterocaulon
clade”

Accepted for publication 23 August 2011

Bean, Pterocaulon section Monenteles

281

Pterocaulon species are readily identified
by the winged stems formed by decurrent
leaf bases, the sessile capitula grouped into
spiciform, cylindrical or spherical secondary
heads (capitulescences), and the pappus of
capillary bristles. In Pterocaulon section
Monenteles , the secondary heads are usually
aggregated to form a globose or ellipsoidal
capitulescence, and only two or three species
have a spicate or cylindrical capitulescence.
Pterocaulon is sometimes confused with the
related genus Sphaeranthus L., as the latter
also has globose capitulescences and winged
stems. However, Sphaeranthus differs by the
much-reduced capitula and the florets that
lack a pappus.

Cabrera & Ragonese (1978) recognised six
species in Pterocaulon section Monenteles ,
including one species newly described
therein. Wilson (1992) reinstated Pterocaulon
globuliflorum W.Fitzg., so that there were
seven accepted species prior to this study,
viz. P. globuliflorum , P. niveum Cabrera &
A.M.Ragonese, P. redolens (Willd.) Fern.-
Vill., P. serrulatum (Montrouz.) Guillaumin,
P. sphacelatum (Labill.) F.Muell.,
P. sphaeranthoides (DC.) F.Muell. and
P. verbascifolium (Benth.) F.Muell.

Fourteen species are recognised in this
paper. The following species are described as
new; Pterocaulon brachyanthum A.R.Bean,
P. ciliosum A.R.Bean, P. discolor A.R.Bean,
P. paradoxum A.R.Bean, P. tricholobum
A.R.Bean and P. xenicum A.R.Bean.
Monenteles intermedins DC. is reinstated
and transferred to Pterocaulon. Pterocaulon
serrulatum is maintained with two varieties.

Materials and methods

This revision is based on morphological
examination of herbarium specimens from A,
AMES, BRI, CANB, CNS, DNA, GH, K, L,
MEL, NOU and PERTH, and images of type
specimens from B, FI, G, K and P. Specimens
from NSW and P were not available for loan
when this paper was being prepared.

All measurements were made from dried
material. Leaf measurements are of cauline
leaves only, and do not include leaves from
the basal rosette (if present). Leaf length is the

distance from the apex to where the midrib
joins the stem. The width of the stem wings
was measured on the main stem at 20-25 cm
from the apex of the specimen.

Measurements of the dimensions of
capitulescences (secondary heads) reported in
this paper were made at or near anthesis, on
capitulescences where the corolla lobes of the
hermaphrodite floret were open, and where
the inner involucral bracts and the florets were
still attached to the receptacle. After anthesis,
and especially in fruiting plants, the inner
bracts and florets become detached from the
receptacle and form expanded “fluffy” heads,
often two or more times the diameter of the
intact heads. Pappus length was measured on
female florets.

Notes on conservation status have
not been given. Most species are common
and widespread; for the species that may
perhaps be threatened (Pterocaulon discolor ,
P. globuliflorum , P. xenicum ), I have no
knowledge of their extent or population sizes.

The catalogue of Malesian plant collectors
(van Steenis-Kruseman 2006-onwards) was
used for checking the interpretation of label
data of Malesian specimens.

Species treatments are arranged in
alphabetical order. Common abbreviations
used in the specimen citations are N.P
(National Park) and S.F. (State Forest).

Chemistry

Species in the genus Pterocaulon are rich in
chemical compounds, particularly coumarins
and flavonoids. These are thought to have a
biocidal effect against bacteria and fungi,
and to protect against herbivory (Daniel et al.
1999).

Johns et al. (1968) identified 6,7-
dimethoxycoumarin as a major alkaloid
constituent of a Pterocaulon from Inkerman,
Queensland, which they identified as
P. sphacelatum. I have located the specimen
on which this is based. It is held at CANB
(Inkerman, 8 June 1960, W.T.Jones 1531),
and it bears a stamp saying “Voucher for
identity of Bulk Sample No. 6489, Aust.
Phytochemical Survey”. The specimen is
Pterocaulon intermedium.

282

MacLeod & Rasmussen (1999) reported
the identification of a new caryophyllene
from Pterocaulon serrulatum, as well
as three coumarins and a flavanone.
Semple et al. (1998) found that a flavonoid
(chrysosplenol C) they extracted from the
leaves of Pterocaulon sphacelatum has an
antiviral effect, by inhibiting the replication
of rhinoviruses, the most frequent causative
agent of the Common Cold. Pterocaulon
species, especially P. sphacelatum and P.
serrulatum have traditionally been used by
Aboriginal peoples as a medicine, helpful in
the treatment of respiratory infections, colds
and fevers (Barr et al. 1993; Latz 1995).

In Asteraceae Subtribe Plucheinae,
Pterocaulon is the only genus known to
contain coumarins, and around 40 different
coumarins have been found in the 15
investigated species (Stein et al. 2007). These
authors stated that although the species
of Pterocaulon exhibit other classes of
compounds, coumarins are characteristic of
the genus and they postulated that they could
be considered chemotaxonomic markers for
the genus.

Ecology and distribution

Taxa belonging to Pterocaulon section
Monenteles are forbs and shrubs. They do not
occur in rainforest or other densely shaded
areas and are primarily found in sunny
shrublands, woodlands and open forests.
Most species are confined to areas where the
climate is frost-free and the rainfall exceeds
c. 600 mm per year. All species show a
preference for coarse-grained soils, probably
because of the associated good drainage, but
can sometimes be found on finer grained clay-
loams. None however, occurs on cracking
clays.

Many species of Pterocaulon clearly
tolerate salt-bearing wind, as a few species are
found in littoral habitats, not only in Australia,
but also in Indonesia and New Caledonia. In
coastal areas, they can be found close to salt
marsh or mangrove communities, and in the
arid areas close to salt lakes. The label for
Henry 952 (NT), collected from the Simpson
Desert, says that Pterocaulon sphacelatum is
“common in saline soil on surrounding edges
of salt lake”.

Austrobaileya 8(3): 280-334 (2011)

It is presumed that Pterocaulons do not
have strict requirements with regard to soil
acidity or soil chemistry. While they frequently
occur on soils derived from sandstone, where
the pH would be quite low, some species
(e.g. Pterocaulon globuliflorum, P. niveum )
occur on alkaline limestone substrates. In
New Caledonia and near Rockhampton in
Queensland, Pterocaulon spp. occur on
serpentinite, and the soils that develop there
are toxic to most plant species.

As with numerous other species in the
Asteraceae, Pterocaulon species are pioneer
plants. Accordingly, they favour places that
have been disturbed by fire or machinery,
and may occur in large numbers at such sites.
For this reason, they are sometimes regarded
as weeds, but they are readily displaced by
alien species that have a similar ecological
preference.

The life cycle is not known in detail
for any species of Pterocaulon section
Monenteles. All species appear to develop a
rootstock of some kind, and in some species
(e.g. P. intermedium ) it is distinctly woody.
Some species are clearly perennial, living for
three or four years at least; while others are
probably annual resprouters (above-ground
parts dying each year, but resprouting for one
or more years).

Pterocaulon section Monenteles is
distributed in southern China, Burma,
Thailand, Indochina, Malesia, much of
Australia (north of about 30°S), New Caledonia
and Vanuatu. The greatest diversity of
species is in the Kimberley region of Western
Australia and the ‘Top End’ of the Northern
Territory. Only one species {Pterocaulon
brachyanthum) does not occur in Australia.

Discussion of characters

1. Winged stems

Winged stems formed from the decurrent leaf
bases are a constant feature of all species in
the genus, and the generic name is derived
from this attribute. The width of the stem
wings is useful in distinguishing some
species in Pterocaulon section Monenteles.
For example, the stem wings of Pterocaulon
niveum are invariably narrow (usually 0.5-

Bean, Pterocaulon section Monenteles

283

1.5 mm wide), while the stem wings of the
vegetatively similar P. verbascifolium are
broad (usually 2.5-6 mm wide). In Pterocaulon
serrulatum and P. sphaeranthoides the stem
wings are serrated; in other species they are
nearly always entire ( P. sphacelatum and
P. brachyanthum sometimes have sparsely
toothed stem wings).

2. Leaves

Some species, and perhaps all species, begin as
a rosette of leaves at ground level. Knowledge
about this stage of the plant’s life cycle is
poor, as these leaves are rarely represented
on herbarium specimens. In some species, the
cauline leaves are obovate or oblanceolate,
i.e. broadest above the middle, while in
others, they are elliptical, i.e. broadest at
around the midpoint. The upper leaf surface
of most species is bullate to a greater or lesser
extent. The leaf margins are always dentate
to denticulate, though sometimes appearing
entire as the teeth can be obscured by dense
indumentum or by recurved margins.

3. Leaf indumentum

Lanate hairs (uniseriate, multicellular)
(Fig. 1A) are found in all species, but their
density varies greatly, with some taxonomic
correlation. In Pterocaulon verbascifolium
and P. niveum , the lanate hairs are dense
enough to obscure the lamina surface even
under magnification; in P. sphaeranthoides
and P. serrulatum var. serrulatum , the lanate
hairs may be very sparse. The upper leaf
surface often has a sparser indumentum
than the lower, and this is particularly the
case in Pterocaulon discolor. In Pterocaulon
intermedium , the lanate hairs on the upper
surface are as dense as or denser than those
on the lower surface.

Two taxa have broad-based stalked
glandular hairs on the leaves and stem wings
(Fig. IB). Globose sessile glands (often
shining-yellow) are found on the lower leaf
surface of all species, and their density can be
diagnostic for some species (Fig. 1A).

4. Capitulescence (secondary head) shape
and diameter

The capitulescences of Pterocaulon
globuliflorum and P. brachyanthum are
consistently narrower in diameter than other
species in the section.

While head shape varies within species,
it is still quite diagnostic. Three species
{Pterocaulon serrulatum, P. redolens and
P. verbascifolium ) have heads usually
described as cylindrical or spicate. In
Pterocaulon paradoxum, capitulescences
are usually markedly ellipsoidal, though
sometimes approaching globose. Similarly,
the heads of Pterocaulon serrulatum are
sometimes almost globose.

For the remainder, heads are globose or
ellipsoidal; in some species (e.g. Pterocaulon
intermedium) the capitulescences are
always close to globose while in others (e.g.
P. sphaeranthoides , P. sphacelatum) they
may vary from globose to broadly-ellipsoidal,
sometimes on the same plant.

5. Branching patterns and peduncles

Capitulescences are in some species
predominantly sessile, while in others
consistently pedunculate. Peduncle length can
be reliably determined only for overtopped
capitulescences, i.e. those where a side branch
has arisen below it, exceeding it and producing
a further (younger) capitulescence. The
peduncle length is the distance between the
base of the overtopped capitulescence and the
insertion of the lateral branchlet. Pterocaulon
intermedium is notable for its consistently
pedunculate heads. The lateral branchlets
usually protrude at 30-60° and sometimes
at right angles to the subtending branch. In
a few species with sessile capitulescences,
e.g. Pterocaulon tricholobum , they may
continue in more-or-less the same direction
(0-30°), here termed ‘continuous’ branchlets
(e.g. Fig. 19). Where this happens, the sessile
capitulescence appears to be borne laterally
and its shape can be globose-truncate.

284

Fig. 1. Leaf hairs in Pterocaulon. A. P. serrulatum var.
velutinum, showing lanate hairs and sessile glands x 40.
B. P. serrulatum var. serrulatum, showing broad-based
glandular trichomes and sessile glands * 40. A from
Bean 22246 (BRI); B from Fox 25 & Bean (BRI). Del.
W. Smith.

6. Inner involucral bracts

The inner involucral bracts offer some
diagnostic characters. Firstly, the bract length
(only the longest (innermost) bracts are
documented in this paper) may be diagnostic
for some species. Their colour may be white,
straw-coloured or pink, or pink with violet
speckles. The colour is not totally fixed for each
species, but some species have predominantly

Austrobaileya 8(3): 280-334 (2011)

pink bracts, some have predominantly white,
and in others the bracts are straw-coloured.

The upper margins of the inner bracts may
be entire, slightly toothed, strongly toothed or
lacerate, and this is strongly correlated to the
species. For example, in Pterocaulon redolens
the upper margins are entire, tapering to an
attenuate apex; in P. sphacelatum, the upper
margins are often lacerate, and there is no
single bract apex.

7. Cilia

The margins of the inner bracts of every
species have ± straight, hyaline, antrorse
cilia, up to 75% of the bract length. The cilia
of different species may vary in number,
position and length.

This character may be usefully employed
on intact or in-situ heads, i.e. where the
capitula are still attached to the receptacle.
The cilia protrude beyond the outer bracts
and may be seen alongside the inner bracts
and the pappus. The cilia of the bracts are
distinguishable from the bristles of the pappus
because the cilia are not barbellate. The
prominence of cilia varies greatly between
species. In some species e.g. Pterocaulon
tricholobum , P. ciliosum , numerous cilia
are clearly visible on the protruding part of
inner bracts, and in P. ciliosum only, these
cilia can exceed the bract apices; in other
species e.g. P. intermedium , P. sphacelatum ,
P. brachyanthum , very few if any cilia can be
seen on the protruding part of inner bracts
(Fig. 2).

8. Number of female florets per capitulum

The number of female florets is difficult
to determine by casual observation of
the capitulum, and an accurate count is
rather tedious, involving removal of an
intact capitulum and spreading out the
contents so that each element is clearly
visible. However, it can be very useful in
distinguishing between some species in
the section. Three groups of species can be
distinguished using this character; Pterocaulon
redolens , P. serrulatum, P. verbascifolium and
P. xenicum with 26-58 florets; P. ciliosum ,
P. globuliflorum,P. paradoxum,P. sphacelatum ,
P. sphaeranthoides and P. tricholobum with

Bean, Pterocaulon section Monenteles

285

Fig. 2. Single capitula in Pterocaulon. A. P. sphacelatum
x 16. B. P. ciliosum x 12. A from Piercey 51 (DNA); B
from Bean 19221 (BRI). Del. W.Smith.

14-29 florets; P. brachyanthum, P. discolor ,
P. intermedium and P. niveum with 7-17
florets.

9. Florets

There is a single hermaphrodite floret in each
capitulum of every species, except Pterocaulon
verbascifolium and P. sphaeranthoides , which
can sometimes have two.

On this floret, a cluster of globose sessile
glands is consistently present on the outer
surface of each corolla lobe, for all species
except Pterocaulon verbascifolium , in which
these glands are lacking. In some species (e.g.
Pterocaulon tricholobum) the corolla lobes
of the hermaphrodite floret consistently bear
numerous conspicuous eglandular unicellular
trichomes on their outer surface. In other
species (e.g. Pterocaulon intermedium) the
lobes are always glabrous. In some species
(e.g. Pterocaulon ciliosum) the trichomes are
absent on some collections, but present on
others.

The length of the corolla lobes of the
hermaphrodite floret is taxonomically
informative. In one group of species, the
corolla lobes are 0.7-1.2 mm long; in another
group, they are 0.3-0.6 mm long. In only
two species (Pterocaulon intermedium and
P. paradoxum) are the lobes of intermediate
length.

The female florets are very slender, with
an inconspicuously lobed corolla and a 2-fid
style (Fig. 20L). The female corolla length
can be diagnostic for species determination.

10. Achenes and pappus

The length of achenes varies from 0.5-1.2 mm
within the section, with each species showing
a small range of lengths. Twin hairs are found
on the achenes of all species, but the number
of hairs varies between species, and is hence
taxonomically useful.

In a few species, it is possible to find a ring
of transparent globose glands at the junction
of the achene and the pappus (Fig. 11L).
These glands are not consistently present in
any species, but can most frequently be seen
in Pterocaulon redolens. The pappus length
varies between species and is often diagnostic,

286

Austrobaileya 8(3): 280-334 (2011)

moz

HERBARIUM BOGORiENSE

Rojor, lodoaeti

Coll.

No.

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HnWali. Calcutta. Kepony, Briitbanc, Canberra. Sandakan, China
Smimnon, For, Res Inst Honor.

Please notify Bog or Herbarium of new identification oF this specimen

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Bean, Pterocaulon section Monenteles

287

but no other pappus characteristics appear to
be of taxonomic utility.

Taxonomy

Pterocaulon section Monenteles (Labill.)
Kuntze, Lex. Gen. Phan. 468 (1903);
Monenteles Labill., Serf. Austro-Caledon. 42,
t. 43, 44 (1825). Type species: Monenteles
spicatus Labill. (= Pterocaulon redolens
(Willd.) Fern.-Vill.,/^ Cabrera & Ragonese
[1978: 202]).

Annually resprouting or perennial forbs or
shrubs, often aromatic. Basal rosette present
in some species. Stems conspicuously
winged due to decurrent leaf margins, wings
extending for 1 and 2 nodes. Leaves alternate,
sessile, decurrent; indumentum of uniseriate,
multicellular hairs; hairs of the upper surface
with short basal cells and relatively long upper
cells; hairs of the lower surface with cell
length relatively uniform throughout; broad-
based glandular hairs sometimes present;
lower surface with yellow, shining, globose
sessile glands. Capitulescences terminal,
monochasially cymose; globose, ellipsoidal,
cylindrical or spiciform; partial clusters
subtended by a bract. Capitula cylindrical,
with involucral bracts in two series; the outer
bracts persistent, lanate-hairy and sometimes
glandular-hairy, all similar in size; the inner
bracts caducous, margins with ± straight,

spreading to antrorse cilia, dorsal surface
glabrous or with tiny translucent sessile
glands, inner whorls slender, outer whorls
shorter and broader. Receptacle epaleate,
glabrous. Outer florets numerous, filiform,
female; corolla three-lobed, style bifid. Disc
floret hermaphrodite, solitary (rarely 2),
tubular, functionally male; corolla broad, 5-
lobed; anthers tailed; style bifid; style branches
with acute sweeping-hairs reaching beyond
the furcation. Achenes narrowly-ellipsoid
to obovoid, with a conspicuous white ring-
shaped carpopodium; twin hairs antrorse,
appressed. Pappus comprising a single row
of barbellate capillary bristles, united at the
base, persistent.

Cytology

The only published cytological record for
Pterocaulon section Monenteles is n = 10 for
Pterocaulon sphacelatum (Watanabe et al.
1999).

Etymology

Monenteles is derived from the Greek
monos meaning ‘single’ and enteles meaning
‘complete, entire or perfect’. This refers to the
single hermaphrodite floret in each capitulum,
surrounded by numerous female florets.

Fourteen species; distributed in Australia,
Melanesia, Malesia and South-east Asia.

Key to the species of Pterocaulon section Monenteles

1 Capitulescences globose-ovoid, spherical to ellipsoidal, 0.7-2.1 times

longer than wide.2

1. Capitulescences cylindrical or spicate, 2.2-5.8 times longer than wide.13

2 Some capitulescences on each plant laterally placed on continuous

branchlets, side branches erect or diverging at less than 30 degrees

(Fig-7).3

2. All capitulescences terminal, side branchlets diverging at 30-90 degrees

(Fig. 5).6

3 Stem wings broad, 2.8-5.5 mm wide .12. P. tricholobum

3. Stem wings narrow, 0.3-2.5 mm wide.4

4 Upper side of leaves with dense to very dense lanate indumentum; inner

involucral bracts pink to violet in dried specimens

6. P. niveum

288 Austrobaileya 8(3): 280-334 (2011)

4. Upper side of leaves with sparse lanate indumentum; inner involucral

bracts greenish-white to white in dried specimens.5

5 Capitulescences 8.5-10 mm wide; pappus 1.3-1.8 mm long .4. P. globuliflorum

5. Capitulescences 11-13 mm wide; pappus 2.6-3.4 mm long .3. P. discolor

6 Stem wings serrate or dentate. 7

6. Stem wings entire or sparsely toothed.8

7 Leaf upper surface conspicuously bullate; few or no cilia visible on

protruding part of inner bracts (Fig. 2A); inner bracts pink, violet or

white.11. P. sphaeranthoides

7. Leaf upper surface not or slightly bullate; many cilia visible on protruding

part of inner bracts (Fig. 2B); inner bracts white or greenish-white . . 9. P. serrulatum

8 Capitulescences 8.5-10 mm wide at anthesis; longest inner bracts 2.8-3.5

mm long; sessile glands on lower leaf surface very sparse.1. P. brachyanthum

8. Capitulescences 10-15 mm wide at anthesis; longest inner bracts 3.4-5.6

mm long; sessile glands on lower leaf surface sparse to dense.9

9 Female florets 8-17 per capitulum; achenes hairs 0-25; leaves elliptic to

lanceolate, broadest near the middle .5. P. intermedium

9. Female florets 16-44 per capitulum; achenes hairs 25-50 or more than

50; leaves oblanceolate to obovate, broadest towards apex.10

10 Corolla lobes of hermaphrodite floret 0.5-0.8 mm long; numerous cilia

visible on protruding part of inner bracts, but none exceeding apices . 7. P. paradoxum

10. Corolla lobes of hermaphrodite floret 0.35-0.6 mm long; cilia on

protruding part of inner bracts either few or absent, or cilia abundant

with some exceeding bract apices.11

11 Stem wings 0.3-0.5 mm wide; longest inner bracts 4.8-5.6 mm long . . . . 14. P. xenicum

11. Stem wings 0.7-3 mm wide; longest inner bracts 3.4-4.6 mm long.12

12 Protrudingpartofinnerbractswithabundantcilia(Fig.2A), some exceeding

bract apices (Fig. 2B); inner involucral bracts straw to white coloured;

leaves 2.1-3.5 times longer than wide.2. P. ciliosum

12. Protruding part of inner bracts with few or no cilia; inner involucral

bracts, white or pink to violet; leaves 3.3-5.8 times longer than wide 10. P. sphacelation

13 Stem wings serrate; achenes 0.7-1 mm long; outer involucral bracts with

broad-based glandular hairs and lanate trichomes outside.9. P. serrulatum

13. Stem wings entire; achenes 0.5-0.7 mm or 1.1-1.3 mm long; outer

involucral bracts with lanate hairs only outside.14

14 Capitulescence 17-30 mm wide at anthesis; hermaphrodite corolla 4-6.5

mm long; achenes 1.1-1.3 mm long.13. P. verbascifolium

14. Capitulescence 8-14 mm wide at anthesis; hermaphrodite corolla 2.5-3

mm long; achenes 0.5-0.7 mm long.8. P. redolens

Bean, Pterocaulon section Monenteles

289

1. Pterocaulon brachyanthum A.R.Bean
species nova affinis P. sphacelato sed
capitulescentiis minoribus, bracteis
interioribus brevioribus, dentibus minoribus
in margine superiore illarum et flosculis
femineis paucioribus in quoque capitulo
differens. Typus: Indonesia. West Nusa
Tenggara Province: Trail from Batudulang
to Punik, Mt. Batulante, west Sumbawa, 25
October 1961, A.J.G.Kostermans 19111 (holo:
BRI; iso: CANB, L).

Shrub 40-70 cm high; stems densely to very
densely lanate; stem wings entire or with
scattered teeth, 0.9-3 mm wide. Leaves
narrowly-elliptic to oblanceolate, 36-55 x
10-22 mm, 2.5-3.6 times longer than wide,
apex acute; margins recurved, denticulate,
with 2-14 pairs of blunt teeth, sometimes
obscure. Indumentum comprising lanate
multi-cellular hairs, glandular hairs absent.
Upper surface with hairs erect to spreading,
sparse to moderately dense, basal cells
short; globose sessile glands absent. Lower
surface with lanate hairs spreading, dense to
very dense, extending 0.4-0.7 mm from the
surface, basal cells not very short; sessile
glands globose, yellow and shining, very
sparse, 8-30 diameters apart. Capitulescence
globose to ellipsoidal, 9—14 mm long and 8.5-
10 mm wide at anthesis, 1-1.5 times longer
than wide; side branchlets never continuous,
at 20-60°; peduncles 2-17 mm long. Outer
involucral bracts 1.5-1.8 mm long, oblong
to oblong-spathulate, with dense spreading
lanate hairs on outer surfaces, inner surface
glabrous or with hairs on the distal end of
inner surface; glands absent; apex obtuse,
erose or truncate. Inner bracts straw-coloured
or white on dorsal surface; longest ones linear,
2.8-3.5 x 0.25-0.5 mm, apex acute, upper
margins with numerous small teeth; cilia
3-9, up to 1.7 mm long, attached along the
proximal half, few extending to the distal one
third, none exceeding the bract apex. Filiform
(female) florets 7-17, colour unknown, corolla
2-2.4 mm long, styles extending 0.3-0.5 mm
beyond corolla. Hermaphrodite floret solitary,
corolla colour unknown, 2.4-2.8 mm long;
corolla lobes 0.3-0.5 mm long, globose sessile
glands present, eglandular trichomes absent.
Achenes narrowly ellipsoidal, 0.5-0.8 mm

long, dark brown, with 0-25 or 25-50 twin
hairs each c. 0.05 mm long; globose sessile
glands at junction between achene and pappus
sometimes present; pappus 2.1-2.5 mm long.

Figs. 3, 4A-D.

Additional specimens examined : Indonesia. Likoeatang
- Taramana, Alor Island, May 1938, Jaag853 (L); Saseel
Island, Kangean Archipelago, Apr 1919, Backer 288635
(L); Karimunjawa Island, north-central Java, Nov 1955,
Hoogerwerf 172 (L); Flores, Lesser Sunda Islands, Sep
1966, Schmutz 355 (L); Soembawa, Nov 1879, Coifs 129
(L); Kambaniroe, Soemba [Sumba] Island, s.dat. [Sep
1873], Teysmann 8824 (A, L); Soemba [Sumba Island],
Lesser Sunda Islands, Nov 1932, Posthumus 3026 (L);
Sumba, Lesser Sunda Islands, Jul 1974, Verheijen 4147
(L); Banyupoh, Bali, Oct 1929, Demandt & VanDillewijn
s.n. (L); Timor, s.dat., Spanoghe s.n. (L); Timor, s.dat.,
Zippelius s.n. (L); Buton Island, south-east Celebes, Sep
1909, Elbert 2598 (L); Timor, Aug 1973, Kooy 957 (K,
L). Timor-Leste, s.loc., 1962-63, Cinatti 210 (L); in
Tasitolu area, just west of Dili, Oct 2006, Cowie 11213
(DNA, L).

Distribution and habitat : Pterocaulon
brachyanthum is endemic to Indonesia and
Timor-Leste. Most collections are from the
Lesser Sunda Islands, east of Java, and from
Timor, but there are two records from islands
north of Java, and one from the south-eastern
end of Sulawesi (Map 1). It inhabits grassy
areas near the coast. One specimen label states
“from the upper margin of mangroves and
salt flat, at the base of a steep shale hilL. The
collections seen were made between altitudes
of 0-200 metres, except the type, which was
from 700 metres.

Phenology : Flowers and fruits are recorded
from April to November.

Affinities : Pterocaulon brachyanthum is
similar to P. sphacelatum, but differs by
the 8.5-10 mm wide capitulescences (10-14
mm wide for P. sphacelatum ); longest inner
bracts 2.8-3.5 mm long (3.5-4.5 mm long for
P. sphacelatum ); upper margins of inner bracts
with small teeth (lacerate or conspicuously
toothed for P. sphacelatum ); female florets
7-17 (17-29 for P. sphacelatum)', and globose
sessile glands very sparse on the lower leaf
surface (dense for P. sphacelatum).

Notes: Cabrera & Ragonese (1978) cited
specimens of the type ( Kostermans 19111 )
from P and NY.

290

Austrobaileya 8(3): 280-334 (2011)

Fig. 4. Pterocaulon brachyantlium. A. hermaphrodite floret x 32. B. inner involucral bract (cilia not shown) x 16.
C. upper portion of inner involucral bract x 32. D. mature achene and basal portion of pappus x 32 . P. ciliosum. E.
hermaphrodite floret x 32. F. inner involucral bract (cilia not shown) x 16. G. upper portion of inner involucral bract
x 32. H. mature achene and basal portion of pappus x 32. P. discolor. I. hermaphrodite floret x 32. J. inner involucral
bract (cilia not shown) x 16. K. upper portion of inner involucral bract x 32. L. mature achene and basal portion of
pappus x 32. A-D from Cowie 11213 (DNA); E,H from Bean 12069 (BRI); F,G from Bean 27963 (BRI); I-L from
Kerrigan 579 (DNA).

Bean, Pterocaulon section Monenteles

291

Distinctive features of this species are
the capitulescences 8.5-10 mm wide; stem
wings often toothed; glands very sparse on
lower leaf surface; female florets few in each
capitulum.

Etymology : From the Greek brachys meaning
short and anthos meaning flower. This is
in reference to both the hermaphrodite and
female florets, which are shorter than in most
other species of the section.

2. Pterocaulon ciliosum A.R.Bean species
nova affinis P. sphacelato sed foliis latioribus,
bracteis interioribus stramineis, ciliis
abundantibus in capitulis in situ visibilibus,
dentibus minoribus in margine superiore in
bracteis interioribus et pedunculis brevioribus
differens. Typus: Australia: Queensland.
Port Curtis District: 7.1 km by road NNW
of Kalpowar Railway Station towards Many
Peaks, 30 August 1975, R.Coveny 6847 &
P.Hind{ holo: BRI; iso: L, NSW).

Illustration : Stanley & Ross (1986: 532), as
P. sphacelatum

Aromatic shrub 40-100 cm high; stems one to
several from non-woody rootstock, sparsely
to densely lanate and sometimes with a few
globose sessile glands; stem wings entire, 1.2-
3 mm wide. Leaves oblanceolate to obovate,
32-56 x 11-21 mm, 2.1-3.5 times longer
than wide, apex acute; margins recurved,
denticulate, with 8-18 pairs of blunt teeth.
Indumentum comprising lanate multi-cellular
hairs, glandular hairs absent. Upper surface
somewhat bullate; hairs erect to spreading,
sparse to moderately dense, basal cells
short; globose sessile glands absent. Lower
surface with lanate hairs erect to spreading,
moderately dense to dense, extending 0.5-1
mm from the surface, basal cells not very
short; globose sessile glands yellow and
shining, dense, touching or up to 3 diameters
apart. Capitulescence globose, 10-16 mm
long and 10-13 mm wide at anthesis, 0.9-
1.2 times longer than wide; side branchlets
never continuous, at 30-90°; peduncles to
7 mm long. Outer involucral bracts 1.8-2.4
mm long, oblong to oblong-spathulate, with
dense spreading lanate hairs on outer surface,
glabrous or with hairs on the distal end of inner
surface; glands absent; apex obtuse. Inner

bracts white to straw coloured (sometimes
tinged pink) on dorsal surface; longest ones
linear, 3.4-4.3 x 0.2-0.4 mm, apex acute,
upper margins with some small teeth; cilia
7-15, up to 2.9 mm long, attached along the
proximal half and consistently extending to
the distal one third, and some exceeding the
bract apex. Filiform (female) florets 16-23,
pink or straw-coloured, corolla 2.2-3 mm
long, styles extending 0.2-0.5 mm beyond
corolla. Hermaphrodite floret solitary, corolla
pink, 2.9-3.5 mm long; corolla lobes 0.35-0.5
mm long, globose sessile glands present,
eglandular trichomes absent or present.
Achenes narrowly ellipsoidal, 0.7-0.9 mm
long, obscurely quadrangular, dark brown,
with 25-50 twin hairs each <0.1 mm long,
evenly distributed; globose sessile glands at
junction between achene and pappus absent;
pappus 2.3-3.1 mm long. Figs. 2B, 4E-H, 5.

Additional selected specimens examined : Indonesia.
Flores, May 1967, Schmutz 1545 (L); Manggarai, Flores,
Jul 1983, Schmutz 5996 (L). Australia: Northern
Territory. Blue Mud Bay, 10 km N of Harris Creek
crossing, Sep 1987, Russeli-Smith 3117 & Lucas (DNA);
Tanumbirini Station, Apr 1991, Wilson 522 (BRI, DNA);
50 miles [80 km] S of Borroloola Road - Anthonys
Lagoon Road junction, Jun 1971, Latz 1539 (DNA); c. 21
km W of ‘Wol logorang’ on the Calvert Hills Road, May
1974, Pullen 9214 (CANB, DNA); 8.7 miles [14.0 km] S of
‘Dunmurra’, Sep 1957, Chippendale 3751 (BRI, CANB,
DNA); Goulburn Island, Anyiminali Point, May 1992,
Coleman 280 (DNA); Caledon Bay, Jun 1972, Latz 2691
(DNA). Queensland. Burke District: 38.5 km NW of
‘Bowthorn’ Homestead on Bowthorn Station, Jun 2006,
Thompson WES745 & Hogan (BRI). Cook District: 4.4
km S of ‘Fairlight’ on the Palmerville Road, Jun 1992,
Clarkson 9607 & Neldner (BRI, CNS, K, MEL); Palm
Cove, Cairns, Jul 1995, Jago 3526 (BRI, DNA); Spear
Creek, c. 1 mile [1.6 km] north of Mt Molloy, Aug 1963,
Schodde 3355 (A, AD, B, BRI, E, L). North Kennedy
District: Hook Island, Whitsunday Group, Jul 1985,
Warrian CW712 (BRI); Mt Abbot, 50 km W of Bowen,
Aug 1992, Bean 4893 (BRI); May’s firebreak, W of
Ravenshoe, Aug 2010, Bean 29858 & McDonald (AD,
BRI, L). South Kennedy District: Brampton Island,
Sep 1988, Batianoff 5114 (BRI); 5.4 km from ‘Redcliffe
Vale’ Homestead on road to Turrawallah, May 1997,
Thompson 266 (BRI). Leichhardt District: Grey Rock,
Glenlea Road, WNW of Springsure, Jul 1989, O ’Keeffe
894 (BRI); 10 km S of Isla Gorge lookout, 37 km S of
Theodore, Jun 1971, Johnson 7169 & Briggs (BRI,
NSW). Port Curtis District: Hill W of Mt Couti-Outi,
Stanage Bay area, N of Rockhampton, Jul 1997, Bean
12069 (BRI); 4 km SW of Mt Castletower, c. 40 km SW of
Gladstone, Jun 1977, Crisp 2739 (BRI, CANB). Burnett
District: Near junction of Delubra and Cadarga Creeks,
35 km SW of Mundubbera, Sep 2008, Bean 27963 (BRI).

292

Austrobaileya 8(3): 280-334 (2011)

Ex- NATIONAL HERBARIUM OF NEW SOUTH WALES

ROYAL BOTANIC GARDENS, SYDNEY

313281

Pterocaulon sphacelatum (Labi11.) F.Muell,

Lot. 24 * 38 'S Long. 151 0 17 ' E

LocJ.lka (4.4mls) by toad NNW of Kalpower
railway atn towards Many Fe-'ks, Old .

Coll. H. Coveny 6847 & P. Bind D °t e 30.Tiii.191

Notes Herb 50 —6 Oct high with mauve flowers,
scattered in sandy creek bed beneath Caauarina
cumin j&amlana . Melaleuca , etc.

Ho£ja~rf P ^ ef C[ueeT ' slan(1 n erbarium ( BR| )

Pfcrocjzuio/L- CthofUmr A. A. &4r\

///^

247327

Brisbane

Fig. 5. Holotype of Pterocaulon ciliosum (Coveny 6847 & Hind [BRI]).

Bean, Pterocaulon section Monenteles

293

Wide Bay District: Fairlies Knob, Seaview Range, May
2000, Phillips 389 & Phillips (BRI); Western edge of
Mt Walsh N.R, near Biggenden, Sep 2002, Bean 19221
(BRI). Moreton District: Upper Ithaca Creek, Aug 1887,
Simmonds s.n. (BRI); Mermaid Mt, NW of Brookfield,
May 1970, Telford 1590 (CANB).

Distribution and habitat: Pterocaulon
ciliosum is a widely distributed species. It
has been collected on the island of Flores
in Indonesia; it occurs in the north-east of
Northern Territory (extending to far north¬
west Queensland), and is widespread in
eastern Queensland from Murray Island in
the Torres Strait to Brisbane (Map 2). It is
found in eucalypt woodland, mainly on well-
drained sandy to loamy soils, including sites
near the ocean.

Phenology : Flowers and fruits have been
recorded from April to October.

Affinities: Pterocaulon ciliosum is similar
to P. sphacelatum , but differs by the leaves
2.1-3.4 times longer than wide (3.3-5 times
for P. sphacelatum ); the straw-coloured
inner bracts on dried material (white or
pink for P. sphacelatum ); abundant cilia
visible on the protruding part of inner bracts
(few or no visible cilia for P. sphacelatum );
the upper margins of the inner bracts with
small teeth (strongly toothed or lacerate for
P. sphacelatum)., and the peduncles to 7 mm
long (to 22 mm long for P. sphacelatum).

Notes: Distinctive features for this species are
the abundant cilia visible on the protruding
part of inner bracts, straw-coloured inner
bracts on dried material and the short corolla
lobes of the hermaphrodite floret.

Etymology: The specific epithet is from the
Latin cilia meaning ‘full of cilia’ and refers
to the numerous cilia on the margins of the
inner bracts.

3. Pterocaulon discolor A.R.Bean species
nova affinis P. tricholobae sed alis caulinis
angustioribus, foliis valde discoloribus,
foliorum dentibus paucioribus, flosculis
femineis paucioribus in capitulo quoque
numeris majoribus pilorum in achenis
differens. Typus: Australia: Northern
Territory. 3 km N of Adelaide River bridge on
Stuart Highway, 3 May 1983, J.D. Briggs 819
(holo: CANB; iso: MEL).

Forb 50-80 cm high, aromatic properties
unknown; stems one to several from woody
rootstock, densely lanate but without globose
sessile glands; stem wings entire, 1-2.5 mm
wide. Leaves elliptical, 35-68 x 13-31 mm,
2-2.7 times longer than wide, apex obtuse or
mucronate; margins not recurved, denticulate
to crenulate, with 10-16 pairs of small teeth.
Indumentum comprising lanate multi-cellular
hairs, glandular hairs absent. Upper surface
not or slightly bullate; hairs erect to spreading,
sparse, basal cell short; globose sessile glands
absent. Lower surface with lanate hairs erect
to spreading, very dense, obscuring surface,
extending 0.2-0.3 mm from the surface, basal
cells not very short; globose sessile glands not
visible. Capitulescence globose to ellipsoidal,
12—15(—19) mm long and 11-13 mm wide at
anthesis, 1—1.2(—1.5) times longer than wide;
side branchlets sometimes continuous, at 0-
60°; peduncles absent or up to 12 mm long.
Outer involucral bracts oblong-spathulate,
2.1-2.4 mm long, with dense spreading lanate
hairs on outer surface, sparse to moderately
dense hairs on the distal end of inner surface,
glands few, apex obtuse. Inner bracts white or
greenish-white on dorsal surface; longest ones
linear, 4.5-5 x 0.3-0.4 mm, apex acute, upper
margins entire or with some small teeth; cilia
8-12, up to 3.3 mm long, attached along the
proximal half and consistently extending to
the distal one third, not exceeding the bract
apex. Filiform (female) florets 7-14, white
or straw-coloured, corolla 27-3.2 mm long,
styles extending 0.3-07 mm beyond corolla.
Hermaphrodite floret solitary, corolla pink,
3.3-4 mm long; corolla lobes 07-1.1 mm long,
globose sessile glands present, eglandular
trichomes 1-5 per lobe. Achenes narrowly
ellipsoidal, 0.8-0.9 mm long, faintly ribbed,
dark brown, with 25-50 twin hairs each
0.1-0.2 mm long, evenly distributed; globose
sessile glands at junction between achene and
pappus absent; pappus 2.6-3.4 mm long. Figs.
4I-L, 6.

Additional specimens examined : Australia: Northern
Territory. Tipperary, May 1963, Muspratt 575 (DNA);
0.5 miles [0.8 km] E of Stuart Highway, 60 miles [98
km] [from Darwin?], Apr 1964, Robinson R343 (DNA);
Nitmiluk N.R, eastern boundary. May 2002, Michell
3875 (DNA); Nitmiluk N.R, Apr 2001, Michell 3329
(DNA); Nitmiluk N.R, Edith Falls, May 2002, Kerrigan

294

Austrobaileya 8(3): 280-334 (2011)

AUSTRALIAN NATIONAL HERBARIUM

mi >iiinii!i ;ihmi mi ii

CBG 8307528

Pt'eroc.aulo^

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FLORA OF AUSTRALIA

HERBARIUM. CANBERRA BOTANIC GARDENS

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Fig. 6. Holotype of Pterocaulon discolor (Briggs 819 [CANB])

Bean, Pterocaulon section Monenteles

295

579 (DNA); Tin Camp Creek, c. 20 miles [32 km] S
of Nabarlek mining camp. May 1973, Hartley 13824
(CANB [2 nd sheet only], DNA, L); 11 km E of Katherine
Gorge ranger station. May 2009, Latz 24367 & Quarmby
(DNA, K, NT).

Distribution and habitat : Pterocaulon
discolor is known from a few scattered
locations in the ‘Top End’ of the Northern
Territory, including Nitmiluk N.P. (Map 3).
It grows on sandstone and quartzite hills and
ridges in mixed open forest or open woodland,
in sandy soil.

Phenology : Flowers and fruits have been
collected in April and May.

Affinities: Pterocaulon discolor is similar
to P. tricholobum, but differs by the stem
wings 1-2.5 mm wide (2.8-5.5 mm for
P. tricholobum ); the lanate indumentum of the
lower leaf surface 0.2-0.3 mm high (0.4-0.8
mm high for P. tricholobum ); the 10-16 pairs
of readily observed leaf teeth (15-30 pairs
of obscure leaf teeth for P. tricholobum)., the
7-14 female florets per capitulum (15-21 for
P. tricholobum ); and the 25-50 hairs on the
achene (0-25 hairs for P. tricholobum).

Notes : This species is distinctive in the
strongly discolorous leaves, few female florets
per capitulum, conspicuous leaf teeth, white or
greenish-white inner bracts and the trichomes
on the corolla lobes of hermaphrodite floret.

Etymology : From the Fatin discolor , meaning
‘of different colours’. This refers to the leaves,
which in this species are strikingly green on
the upper surface and snowy white on the
lower surface.

4. Pterocaulon globuliflorum W.Fitzg., J.
& Proc. Roy. Soc. Western Australia 3: 223
(1918), (as “ globuliflorus ”). Type: Australia:
Western Australia. Devil’s Pass, Napier
Range, May 1905, W.VFitzgerald 607 (holo:
PERTH; iso: K373285; NSW n.v).

Shrub to 60 cm high; aromatic properties
unknown; stems sparsely to densely lanate and
with a few globose sessile glands; stem wings
entire, 0.3-1.5 mm wide. Feaves elliptical,
18-51 x 7-24 mm, 2-2.6 times longer than
wide, apex acute or obtuse; margins not
recurved, denticulate, with 7-15 pairs of blunt
teeth. Indumentum comprising lanate multi¬

cellular hairs, glandular hairs absent. Upper
surface not or slightly bullate, hairs sparse,
flexuose, basal cells not short; globose sessile
glands absent. Fower surface with lanate
hairs flexuose, dense to very dense, extending
0.15-0.25 mm from the surface, basal cells not
very short; globose sessile glands yellow and
shining, dense, touching or up to 3 diameters
apart, but usually obscured by indumentum.
Capitulescence globose or globose-truncate,
6—10 mm long, 8.5-10 mm wide at anthesis,
0.7-1.1 times longer than broad; sidebranchlets
0-50°, often continuous, peduncles 0-4 mm
long. Outer involucral bracts oblong, 0.8-1.9
mm long, with dense spreading lanate hairs
on the distal one-half of outer surface, and
glabrous on the inner surface; apex obtuse or
truncate. Inner bracts white on dorsal surface,
sessile glands absent; longest ones linear, 3.1-
3.5 x 0.2-0.25 mm, apex acute, upper margins
with small teeth; cilia 5-8, up to 1.6 mm long,
mostly attached proximally and extending to
the distal one third. Filiform (female) florets
16-20, colour unknown, corolla 1.7-2 mm
long, styles extending 0.3-0.5 mm beyond
corolla. Hermaphrodite floret solitary, corolla
pink, 2.5-2.8 mm long; corolla lobes 0.7-1.2
mm long, globose sessile glands present,
eglandular trichomes absent or with 1 or 2 per
lobe. Achenes narrowly ellipsoidal, 0.7-0.8
mm long, slightly longitudinally ribbed, dark
brown, with 25-50 twin hairs each c. 0.05
mm long, evenly distributed; globose sessile
glands at junction between achene and pappus
absent; pappus 1.3-1.8 mm long. Figs. 7, 8A-
D.

Additional specimens examined : Australia: Western
Australia. Windjana Gorge, Napier Range, Jul 1974,
Carr 3882 & Beanglehole 47660 (PERTH); Windjana
Gorge, Napier Range, Jul 1974, Carr 4036 & Beanglehole
47814 (PERTH); Yammera Gap, Napier Range, 9 km NE
of Lennard River crossing on Gibb River Road, May
1988, Streimann 8269 (A, B, CANB, L); Old Fossil
Downs Station, May 1970, Wolfe & Martin 72 (CANB).

Distribution and habitat : Endemic to the
Napier and Geikie Ranges, Kimberley region,
Western Australia (Map 3). It is confined
to limestone ranges and hills, growing in
grassland or low open woodland.

Phenology: Flowers and fruits have been
recorded in May and July.

296

Austrobaileya 8(3): 280-334 (2011)

Flora of Australia

AUSTRALIAN NATIONAL BOTANIC GARDENS
HERBARIUM (CBG)

Notificatjon of change of determination would be appredated by CBG

AMARANTHACEAE

AUSTRALIA. WESTERN AUSTRALIA. Yammera Gap.
Napier Range, 9 Km NE of Lennard River crossing on Gibb
River Road.

c6A6

PUfocavalon ntoeuifn C a W<

Queensland Herbarium (BRI)

j/oSaJ/f/ort*, &

Det 20 //

17° 20' S 124" SITE 70 m alt.

Grasslands at base of limestone range with scattered small
shrubs.

Open spreading herb to 40 cm tall. Inflorescence white;
leaves dull dark green above, grey-green below.

H. Strelmann 8269 IS May. 1988

Dupl.t (6) L, DIVA, A. PERTH. l^E. B

Fig. 7. Representative specimen of Pterocaulon globuliflorum (Streimann 8269 [CANB]).

Bean, Pterocaulon section Monenteles 297

Fig. 8. Pterocaulon globuliflorum. A. hermaphrodite floret * 32. B. inner involucral bract (cilia not shown) x 16. C.
upper portion of inner involucral bract x 32. D. mature achene and basal portion of pappus x 32. P. intermedium. E.
hermaphrodite floret x 32. F. inner involucral bract (cilia shown) x 16. G. upper portion of inner involucral bract x 32.
H. mature achene and basal portion of pappus x 32. A-D from Carr 4036 & Beauglehole 47814 (PERTH); E-G from
Bean 12186 (BRI); H from Clarkson 4854 (BRI).

Affinities : Pterocaulon globuliflorum appears
to be allied to P. tricholobum and P. niveum ,
on the basis of the long corolla lobes of
the hermaphrodite floret, and the often
‘continuous’ branchlets giving rise to lateral
capitulescences.

Typification: Fitzgerald wrote the location
‘Devil’s Pass’ on the label of his specimen. In
the protologue, he used the locality ‘Wingrah
Pass’. According to an annotation by Annette

Wilson on the holotype, both of these terms
are equivalent to Windjana Pass or Windjana
Gorge.

Notes : Distinctive features for this species
include the small capitulescences (8.5-10 mm
diameter) that are often lateral; leaves short
and broad, 2-2.6 times longer than wide,
strongly discolorous; stem wings very narrow
and the very short pappus.

298

5. Pterocaulon intermedium (DC.) A.R.Bean
comb, nov.; Monenteles intermedins DC.,
Prodr. 5: 456 (1836). Type: [Australia:
Queensland. North Kennedy:] Sandy shores of
Cape Cleveland, 14 June 1819 , A.Cunningham
52 (holo: G-DC [G214737]; iso: K373284).

Monenteles globiferus DC., Prodr. 5: 455
(1836). Type: [Australia: Queensland.
Moreton District:] Lockyer’s Creek, Brisbane
River, Moreton Bay, July 1829, A. Cunningham
51 (holo: G-DC [G214749]; iso: K373282).

Pterocaulon sp. A; Wilson (1992: 954).

Pterocaulon sp. A Kimberley Flora (B.J.Carter
599); Paczkowska & Chapman (2000: 172).

Pterocaulon sp. (Yarrowmere Station E.J.
Thompson+ BUC340); Bostock & Holland
(2007: 30); Bostock & Holland (2010: 25).

Illustration : Kenneally et al. (1996: 67), as
‘Pterocaulon sp. A, Kimb. Flora’

Aromatic perennial shrub 20-60 cm high;
stems sprouting from a woody rootstock,
sparsely to densely lanate and sometimes
with a few globose sessile glands; stem wings
entire, 0.2-1.4 mm wide. Leaves lanceolate,
narrowly-elliptic or elliptic, 12-52 x 3.5-
18 mm, 2.6-4.6 times longer than wide,
apex acute; margins not recurved, entire or
denticulate, with 2-20 pairs of blunt teeth.
Indumentum comprising lanate multi-cellular
hairs, glandular hairs absent. Upper surface
not bullate, hairs moderately dense to very
dense, erect near base then antrorse, basal cells
short; globose sessile glands absent. Lower
surface with lanate hairs erect to spreading,
moderately dense to dense, extending 0.5-1
mm from the surface, basal cells not very
short; globose sessile glands yellow and
shining, dense, touching or up to 3 diameters
apart. Capitulescence globose or almost so,
10-15 mm long, 10-14 mm wide at anthesis,
1-1.2 times longer than broad; side branchlets
never continuous, at 30-90°; peduncles (5-)7-
25 mm long. Outer involucral bracts oblong
to spathulate, 1.7-2.4 mm long, with dense
spreading lanate hairs on the distal one-third
of outer surface, and glabrous or with a few
hairs distally on the inner surface, glands
absent; apex acute, obtuse or truncate. Inner

Austrobaileya 8(3): 280-334 (2011)

bracts pink to violet on dorsal surface; longest
ones linear, 4-4.8 x 0.2-0.4 mm, apex acute,
upper margins with small teeth; cilia 1-7, up
to 2.8 mm long, mostly attached proximally
and not extending to the distal one third.
Filiform (female) florets 8-17, pink, corolla
27-3.6 mm long, styles extending 0.3-0.8
mm beyond corolla. Hermaphrodite floret
solitary, corolla pink, 3.1-3.8 mm long; corolla
lobes 0.6-0.9 mm long, globose sessile glands
present, eglandulartrichomes absent. Achenes
narrowly ellipsoidal, 0.6-0.75 mm long,
longitudinally ribbed, dark brown, glabrous
or with 1-25 twin hairs each <0.1 mm long,
evenly distributed; globose sessile glands at
junction between achene and pappus absent;
pappus 2.5-3.1 mm long. Figs. 8E-H, 9.

Additional selected specimens examined : Indonesia.
Kisar Island [just N of Timor], Apr 1939, Bloembergen
3852 (A, L). Papua New Guinea. Morobe Province:
Sialum Station, May 1975, Henty & Katik NGF49770
(BRI, CANB, L). Australia: Western Australia. Fence
line at junction of Pindan, Calanjadie and Cardingy
paddocks, Anna Plains Homestead, Sep 2004, Byrne 1266
(PERTH); Sir Graham Moore Island, Jul 1973, Wilson
11263 (PERTH); adjacent to Telecom tower, c. 5 km E
of Broome, Oct 1993, Dureau DD150 (BRI, PERTH);

4 km SSE of Cape Bertholet, Dampier Peninsula, Aug
1992, Carter BJC599 (PERTH); Pender Bay at Weedong
Lake, Dampier Peninsula, Jun 1984, Forbes 2435 &
Kenneally (L, MEL). Northern Territory. Elcho Island,
Warangaya, Sep 1987, Russell-Smith 3282 & Lucas (BRI,
DNA); Elcho Island, Refuge Bay, Aug 1995, Cowie 6002
(BRI, CANB, DNA, MEL); Howard Island, Jun 1996,
Booth 1789 (DNA); Maria Island, Gulf of Carpentaria,
Jul 1972, Dunlop 2803 (BRI, DNA); Homestead Creek,
Bing Bong Station, Jun 1971, Dunlop 2245 (DNA).
Queensland. Burke District: Karumba, Jun 1901,
Bailey s.n. (BRI [AQ270420]); Pandanus Ridge, 20 km
NE of Burketown, May 2005, Booth 4281 & Thompson
(BRI); Settlement Creek, 2 miles [3 km] from coast.
Gulf of Carpentaria, Jun 1948, Perry 1217 (BRI, CANB,
DNA); Sweers Island, South Wellesley group. Gulf of
Carpentaria, Nov 2002, Thomas SWI111 (BRI). Cook
District: Lakefield N.P, Jane Table Hill, Jun 1983,
Clarkson 4854 (BRI); near California Creek, NW of Mt
Garnet, Aug 1997, Bean 12186 (BRI); 1.5 km N of the
North Kennedy River on the Laura - Musgrave Road,
May 1989, Clarkson 7996 & Neldner (BRI, DNA, K,
L); on southern bank of Nassau River 0.5 km upstream
from junction with Rocky Creek, Jun 1990, Neldner
3001 (BRI, L). North Kennedy District: 15 miles [25
km] NW of Bowen towards Townsville, May 1970, Fagg
698 (CANB, L). South Kennedy District: 22.5 km

5 of ‘Yarrowmere’ Homestead, Apr 1992, Thompson
BUC340 & Simon (AD, BRI). Port Curtis District:
‘Galloway Plains’, c. 28 km SW of Calliope, Mar 1989,
Anderson 4623 (BRI); S.F.60 Rundle Range, 37 km NW
of Gladstone, Aug 1989, Gibson TOI683 (BRI). Burnett

Bean, Pterocaulon section Monenteles

299

yueensiana neroanum idkij

QUEENSLAND HERBARIUM (BRI)

Fkxa of Queensland Burt<e

Pterocaulon sphacelatum (Labill.) F.Muell.

Pfetioctud*^ A A, Scan

Dei - A*. 6e^ Dale to 61 2 * 1 ,

QUEENSLAND HERBARIUM (BRI)
Brisbane Australia

7323

Fig. 9. Representative specimen of Pterocaulon intermedium (Booth 4281 & Thompson [BRI]).

300

District: Brian Pastures near Gayndah, Mar 1952, Blake
18894 (BRI); Wetheron, Apr 1980, Forster PIF739 (BRI).
Darling Downs District: ‘Rockwood’, 60 miles [98 km]
W of Dalby, Apr 1969, Dunlop 337 (CANB). Moreton
District: Ipswich, 1909, Hall s.n. (BRI [AQ270443]).

Distribution and habitat : Pterocaulon
intermedium is widely distributed in northern
Australia from Port Hedland (Western
Australia) to Ipswich (Queensland). It is often
found close to the coast, but it may occur up to
250 km inland. Outside Australia, it is known
from the island of Kisar in Indonesia, and
from one location in Papua New Guinea (Map
4). In coastal areas it favours sandy swales or
dunes, in shrubland or low woodland, and is
sometimes adjacent to saline tidal areas. In
the more inland areas, it grows in eucalypt
woodland with sandy soil.

Phenology: Flowers and fruits are recorded
from April to October.

Affinities: Pterocaulon intermedium is
perhaps closest to P. paradoxum, but differs
by the leaves being broadest near the midpoint,
the upper surface not bullate, the indumentum
of the upper surface as dense as or denser than
the lower, the shorter achenes and the fewer
female florets per capitulum.

Typification: The original label of the
holotype of Monenteles intermedius gives the
date of collection as “June 14 1829”. This is
incorrect. The isotype at K bears the correct
date of 14 June 1819, as confirmed by Curry
& Maslin (1990).

Notes: Distinctive features for Pterocaulon
intermedium are the narrow stem-wings;
consistently long peduncles; a consistently
± globose capitulescence, inner bracts pink
or violet; relatively few female florets per
capitulum; elliptical leaves; upper leaf surface
indumentum as dense as or denser than lower
surface; and few hairs on the achene.

One specimen label reports that Pterocaulon
intermedium has an “apple juice smell”.

6. Pterocaulon niveum Cabrera &
A.M.Ragonese, Darwiniana 21: 249 (1978).
Type: Australia: Western Australia. Mt
Amherst, 13 May 1951, C.A.Gardner 10206
(holo: PERTH; iso: SI).

Austrobaileya 8(3): 280-334 (2011)

Illustration: Cabrera & Ragonese (1978:
248).

Spreading shrub 40-100 cm high; stems very
densely lanate; stem wings entire, 0.5-1.5
(-2.5) mm wide. Leaves elliptical or obovate,
39-70 x 21-38 mm, 2.1-2.6 times longer
than wide, apex acute or obtuse; margins not
recurved, denticulate, with 9-20 pairs of blunt
teeth. Indumentum comprising lanate multi¬
cellular hairs, glandular hairs absent. Upper
surface with hairs erect to spreading, very
dense; globose sessile glands absent. Lower
surface with lanate hairs erect to spreading,
very dense, extending 0.5-1 mm from the
surface; globose sessile glands sparsely
present, but hidden by hairs. Capitulescence
globose to globose-truncate, 10-15 mm long
and 10-15 mm wide at anthesis, 0.7-1.1 times
longer than wide; side branchlets sometimes
continuous, at 0-70°; peduncles 0-8 mm
long. Outer involucral bracts 2.2-3.2 mm
long, oblong to oblong-spathulate, with dense
spreading lanate hairs on outer surface, inner
surface glabrous; glands absent; apex acute or
obtuse. Inner bracts pink to violet on dorsal
surface; longest ones linear, 4.5-5.4 x 0.2-
0.35 mm, apex acute; upper margins entire or
with a few small teeth; cilia 8-14, up to 3 mm
long, attached mainly along the proximal half
and consistently extending to the distal one
third, not exceeding the bract apex. Filiform
(female) florets 11-17, pink, corolla 3-3.7 mm
long, styles extending 0.4-1.2 mm beyond
corolla. Hermaphrodite floret solitary, corolla
pink, 3.3-4 mm long; corolla lobes 0.7-1.1
mm long, globose sessile glands present,
eglandular trichomes present, 2-8 per lobe.
Achenes narrowly ellipsoidal, 0.9-1 mm long,
terete or slightly angular, dark brown, with
25-50 or >50 twin hairs each c. 0.1 mm long,
evenly distributed; globose sessile glands at
junction between achene and pappus absent;
pappus 2-3.1 mm long. Figs. 10,11A-D.

Additional specimens examined: Australia: Western
Australia. 19 miles [31 km] NW of Turner River Station,
Jul 1949, Perry & Lazarides 2410 (CANB, DNA); China
Wall, Jun 1983, MacDonald 234 (BRI); Mt Amherst
Range, 1 mile [1.6 km] E of Mt Amherst Station, July
1959, Lazarides 6341 (BRI); 9 km SSW of Bungle
Bungle outcamp. East Kimberley, Jul 1984, Kenneally
9264 (BRI, CANB, PERTH); S slopes of Mt North, May
1988, Goble-Garratt 525 (BRI, PERTH); Mt House

Bean, Pterocaulon section Monenteles

301

Spreading, dense sub-shrub 3-4 ft high with
purple, globular flower*. Stems and leaves
woolly and white.

Thi* specimen is presented on the condition that new
identifications will be communicated to CS.T.R.O,, Canberra.

Coll. M. Lazarides 6341 Date 15 July, 1959

Loc. Mt. Amherst Range, 1 mile E. of Mt. Amherst
Station, Kimberley*, W.A.

Occasional on rocky granitic slopes with Triodia spp.

139800

270496

EX HERBARIO A USTR ALIENS!

CS.I.RXX/ Canboira

Collected and distributed by Division of Land Research and
Regional Survey, GSJ.R.O., Canberra

Name Pterocaulon sphaceLatum (Lablll.)

Be nth. et Hook.

M. Lazarides 30 June, i960

det - L LL

Ai l.yq Herb. gR/

Fig. 10. Representative specimen of Pterocaulon niveum {Lazarides 6341 [BRI]).

302

Austrobaileya 8(3): 280-334 (2011)

Fig. 11. Pterocaulon niveum. A. hermaphrodite floret x 32. B. inner involucral bract (cilia not shown) x 16. C.
upper portion of inner involucral bract x 32. D. mature achene and basal portion of pappus x 32. P. paradoxum. E.
hermaphrodite floret x 32. F. inner involucral bract (cilia not shown) x 16. G. upper portion of inner involucral bract
x 32. H. mature achene and basal portion of pappus x 32. P. redolens. I. hermaphrodite floret x 32. J. inner involucral
bract (cilia not shown) x 16. K. upper portion of inner involucral bract x 32. L. mature achene and basal portion of
pappus x 32. A-D from Brocklehurst 372 (DNA); E-G from Keighery 9018 (PERTH); H from George 13582 (PERTH);
I-K from Rodd & Hardie 4378 (BRI); L from McKee 10198 (CANB).

Bean, Pterocaulon section Monenteles

303

Station, Aug 1950, Royce 3304 (PERTH); Caroline Pool,
8 km E of Halls Creek, Jun 1976, Beauglehole ACB53312
(PERTH); 48.5 km N of ‘Springvale’ Homestead, 86
km N of Halls Creek, Jun 1976, Beauglehole ACB53527
(BRI, PERTH); Palm Springs, S of Halls Creek, Aug
2000, Handasyde & Start TH00287 (PERTH); McSpeery
Gap, Napier Range, Jun 1971, Maconochie 1298 (BRI,
DNA); between Njitparriya and Djimbitjba, 3 km SE of
Bungle Bungle Outcamp, Jul 1984, Forbes 2597 (BRI,
CANB, DNA, MEL, PERTH); Mt Broome, May 1971,
Byrnes 2253 (K). Northern Territory. Calcite flow stop.
Limestone Gorge, Gregory N.P, Jun 2000, Kerrigan &
Risler 171 (DNA); Mt Napier area. May 1974, Dunlop
4066 (DNA); Gregory N.P, Fire plot 41, May 1999,
Michell & Johnson 2449 (DNA, NT); Gregory N.P,
28 km SW of Bullita outstation, Apr 1996, Coles 28
& Barritt (DNA, MEL); Gregory N.P, Humbert River
road, Aug 1991, Brocklehurst 372 (DNA).

Distribution and habitat: Pterocaulon
niveum is widespread in the Kimberley region
of Western Australia, occurring in the Napier
Range (western Kimberley) and from Mt
Amherst (south-eastern Kimberley) eastwards
to the Gregory N.R in the Northern Territory
(Map 2). It grows in Triodia grassland or
low open eucalypt woodland, on rocky
slopes or in gullies with shallow sandy soils.
The geological substrate may be sandstone,
quartzite or limestone.

Phenology: Flowers and fruits have been
collected from April to August.

Affinities: Pterocaulon niveum, P.

tricholobum, P. discolor and P. globuliflorum
appear to be related, based on similarity in
morphology of the hermaphrodite floret,
i.e. corolla lobes long (07-1.2 mm long)
with eglandular trichomes present, and the
presence of lateral capitulescences.

Sterile specimens of Pterocaulon niveum
and P. verbascifolium may be confused, as
they have leaves of similar size, shape and
toothing, and the indumentum is extremely
dense for both. However, the much broader
stem wings of Pterocaulon verbascifolium
allow them to be easily distinguished.

Notes: Distinctive features of Pterocaulon
niveum are the indumentum of the lower leaf
surface being extremely dense and obscuring
the surface, the continuous branchlets with
lateral capitulescences present, narrow
stem wings and long corolla lobes on the
hermaphrodite floret.

7. Pterocaulon paradoxum A.R.Bean
species nova affinis P. sphacelato sed alis
caulinis latioribus, glandibus sparsius in
pagina inferiore folii distributis, dentibus
parvis in marginibus superioribus bractearum
interiorum, capitulescentiarum plus
elongatarum et lobis longioribus corollae
flosculi hermaphroditi differens. Typus:
Australia: Western Australia. 15 km ESE of
Winnama yards by Samin mining exploration
track, c. 8 km E of Palms yard, SE Kimberley,
15 May 1984, S.J.Forbes 2017 (holo: DNA;
iso: CANB, MEL).

Illustration: Kenneally et al. (1996), as
P. sphacelatum.

Shrub 60-120 cm high, aromatic properties
unknown; stems one or more from non-woody
rootstock, sparsely to densely lanate but
without globose sessile glands; stem wings
entire, 1.5-4.5 mm wide. Leaves elliptical
to oblanceolate, 30-65 x 10-20 mm, 2.6-4.6
times longer than wide, apex obtuse; margins
recurved, denticulate, with 13-23 pairs of very
small teeth. Indumentum comprising lanate
multi-cellular hairs, glandular hairs absent.
Upper surface slightly bullate; hairs erect to
spreading, moderate to very dense, basal cell
short; globose sessile glands absent. Lower
surface with lanate hairs erect to spreading,
dense to very dense, extending 0.4-0.7 mm
from the surface, basal cells not very short;
globose sessile glands yellow and shining,
moderately dense, 3-6 diameters apart.
Capitulescence ovoid to ellipsoidal, 14-28
mm long and 12-15 mm wide at anthesis, 1.2-
2.1 times longer than wide; side branchlets
never continuous, at 20-50°; peduncles 0-20
mm long. Outer involucral bracts spathulate,
2.4-27 mm long, with dense spreading lanate
hairs on outer surface, sparse hairs on the
distal end of inner surface; glands absent;
apex obtuse or truncate. Inner bracts white to
greenish (occasionally tinged pink) on dorsal
surface; longest ones linear, 3.5-4.8 mm long
and 0.2-0.4 mm wide, apex acute, upper
margins entire or with some small teeth; cilia
6-12, up to 3.1 mm long, attached along the
proximal half and consistently extending to
the distal one third, not exceeding the bract
apex. Filiform (female) florets 18-24, pink,

304

Austrobaileya 8(3): 280-334 (2011)

Lat. 17°14 1 S Long. 128°20*E Alt.: 380 m
Coll.: S-J. Forbes, no.2017, 15.v. 1984
Freq.: occasional

Dei.:

Notes: sandy flat with Eucalyptus confertiflora
Crotalaria novae-hollandiae , Heteropogon
contortus , Lysiphyllum cunninghamii
Habit; erect herb
Flowers white

-Notes: winged stems, weakly aromatic

HoioTyfiE' Queensland Herbarium (BRI)

Pferoca u/on pa,ra.i{e>xet/n d.A.8ca.n

Compositae

* A *4-fci20//

NATIONAL HERBARIUM OF VICTORIA (MEL)

MELBOURNE, AUSTRALIA

Pterocaulon sphacelatum (Labill.) Benth &

Hook.f» ex F.Muell.

Loc.: W.A, - SE Kimberley, (15 km ESE Winnama
yards by Samim mining exploration track; c. 8 kir
E Palms Yard

Fig. 12. Holotype of Pterocaulon paradoxum {Forbes 2017 [DNA]).

Bean, Pterocaulon section Monenteles

305

corolla 2.5-3.4 mm long, styles extending
0.3-07 mm beyond corolla. Hermaphrodite
floret solitary, corolla pink, 2.9-37 mm
long; corolla lobes 0.5-0.8 mm long, globose
sessile glands present, eglandular trichomes
absent. Achenes narrowly ellipsoidal, 0.9-
1.2 mm long, obscurely quadrangular, dark
brown, with 25-50 or >50 twin hairs each
c. 0.1 mm long, evenly distributed; globose
sessile glands at junction between achene and
pappus occasionally present; pappus 2.5-3.5
mm long. Figs. 11E-H, 12.

Additional selected specimens examined : Papua New
Guinea. Central Province: Rogers Airstrip, c. 8 miles
[13 km] W of Kanosia Plantation, Kairuku subdistrict,
July 1962, Darbyshire 647 (A, BRI, CANB, L). Australia:
Western Australia. Adjacent to King Edward River, 8.7
km NW of Gibb River - Kalumburu road intersection,
along old Mitchell River Station Road, Jun 1987, Koch
506 (PERTH); S side of Cockburn Range, c. 13 km W
of King River, Jul 1974, Carr 3338 & Beauglehole 47116
(CANB); King Edward River, 100 km S of Kalumburu,
Jun 1987, Keighery 9018 (PERTH); King Edward River,
Mitchell Plateau road, c. 200 km W of Wyndham, Jun
1976, Beauglehole ACB51933 (PERTH); ‘Beverley
Springs’ Homestead, May 1979, Muir et al. 757
(PERTH); near Dromaius Creek, near southern end of
Ashton Range, Drysdale River N.P, Aug 1975, George
13191 (PERTH); Mogurnda Creek, near Drysdale River,
Drysdale River N.P, Aug 1975, George 13582 (PERTH);
upper reaches of Barker River, 2 km N of ‘Mount Hart’
Homestead, Jun 1987, Edinger 423 (BRI, PERTH);
Oobraguma Road, 5 km E of Stuart River, 74 km NNE
of Derby, Jun 1976, Beauglehole ACB52918 (PERTH);
c. 100 m W of Taylor’s Lagoon, Sep 2005, Byrne 1639
(PERTH); One Arm Point, Dampier Peninsula, Jul 1989,
Carter 410 (PERTH); Mornington Station, Trapline
1, May 2004, Handasyde 2099 (PERTH). Northern
Territory. Middle Beach swamp. West Alligator Head,
Jul 1990, Brennan Bre646 (DNA); Near Maningrida,
Anamayirra Creek, Aug 1995, Cowie 5909 (CANB,
DNA, MEL); Danger Point, Coburg Peninsula, Jul 1961,
Chippendale 8244 (BRI, CANB, DNA); Grant Island,
north side. May 1987, Clark s.n. (DNA); Ramingining
area, W of Dhabla Road, Aug 1998, Cowie & Dunlop
7922 (DNA); Wunyu Beach in Aurari Bay, c. 30 km SE
of Murgenella, Jun 1988, Weber 10102 (AD, DNA).

Distribution and habitat: Pterocaulon
paradoxum is relatively widespread in the
Kimberley region of Western Australia, is
scattered along the north coast of Northern
Territory, and there is one record from Papua
New Guinea (Map 5). It inhabits eucalypt
woodland on alluvial flats (species including
Eucalyptus miniata A.Cunn. ex Schauer,
E. tetrodonta F. MuQ\\.,Corymbiaconfertiflora
(F.Muell.) K.D.Hill & L.A.S.Johnson), on

sheltered sandstone scree slopes, or on coastal
dunes. Soils include red-brown clayey sand,
red loam, and grey sand.

Phenology: Flowers and fruits are recorded
from April to September.

Affinities: Pterocaulon paradoxum appears
to be most closely allied to P. sphacelatum.
It differs by the stem wings 1.5-4.5 mm
wide (0.7-2 mm wide for P. sphacelatum).,
the corolla lobes of the hermaphrodite floret
(0.5-)0.6-0.8 mm long (0.4-0.6 mm long for
P. sphacelatum ); the globose sessile glands
on the lower leaf surface moderately dense,
3-8 diameters apart (dense, 0-3 diameters
apart for P. sphacelatum ); the upper margins
of the inner bracts with small teeth (strongly
toothed or lacerate for P. sphacelatum ); and
the capitulescences usually more elongated.

The capitulescences of this species
provide a link between the ‘spicate’ group
{Pterocaulon redolens, P. verbascifolium ,
P. serrulatum ) and the ‘globose to
ovoid’ group (P. sphacelatum, P. ciliosum,
P. sphaeranthoides, P. intermedium). The
capitulescences are more elongated that other
species in the ‘globose to ovoid’ group, but
not clearly cylindrical as in the ‘spicate’
group. One or more separate clusters of
capitula can usually been seen at the base
of the capitulescence, as in the spicate
species Pterocaulon redolens. In some areas,
particularly the Dampier Peninsula (Western
Australia), the capitulescences appear
consistently almost globose, although the
floret dimensions and other features appear to
be the same as in the type.

Notes: Distinctive features of Pterocaulon
paradoxum are the capitulescences often
markedly ellipsoidal, the medium length
corolla lobes on the hermaphrodite floret and
the long achenes.

Two forms of this species exist. The typical
form is endemic to the Kimberley, mainly
away from the coast, and is a densely hairy
plant with corolla lobes on the hermaphrodite
floret 0.6-0.8 mm long. The second form (e.g.
Cowie 5909) is found along the northern coast
of the Northern Territory, often in littoral
areas, and one specimen from Papua New

306

Guinea is rather doubtfully included here.
The indumentum on vegetative parts is much
sparser in this form than the typical one, there
are few cilia visible on protruding part of inner
bracts, the corolla lobes on the hermaphrodite
floret are shorter (0.5-0.6 mm long) and the
capitulescences are narrower. Both forms
can have ellipsoidal capitulescences, where
separate clusters of capitula are discernable.

Etymology : From the Latin paradoxus ,
meaning ‘contrary to expectations,
puzzling’. This species is puzzling because
of its sometimes ± globular and sometimes
markedly ellipsoidal capitulescences, and the
different forms within the species.

8. Pterocaulon redolens (Willd.) Fern.-Vill.,
Nov. App. ed. 3, 4(3): 116 (1880); Conyza
redolens Willd., Sp. PI. ed. 3(3), 1951 (1804);
Tessaria redolens (Willd.) Less., Linnaea 6:
151 (1831); Monenteles forsteri Endl., Ann.
Wiener Mus. Naturgesch. 1: 168 (1836), nom.
illeg:, Monenteles redolens (Willd.) DC.,
Prodr. 5: 455 (1836). Type: New Caledonia, in
1774, J.R.Forster & J.G.A.Forster (holo: B-W
No. 15639; iso: K373288, 1 st , 2 nd and 4 th pieces
from the left).

Monenteles spicatus Labill., Sert. Austro-
Caledon. 43 (1825); Pterocaulon billardierei
F.Muell., Descr. Notes Papuan PI. 8: 43 (1886),
nom. nov.; Pterocaulon spicatum (Labill.)
Domin, Biblioth. Bot. 89: 664 (1930), nom.
illeg. non DC. (1836). Type: New Caledonia.
[June 1792], J.J.H. Labillardiere (lecto: FI
[FI-W92604], here designated; isolecto: FI
[FI-W92605], right hand specimen only, FI
[FI-W92611], P537803).

Sphaeranthus elongatus Blanco, FI. Filip. 636
(1837), fide Merrill (1918: 379).

Pterocaulon cylindrostachyum C.B.Clarke,
Compos. Ind. 98 (1878), nom. illeg. {Monenteles
spicatus is listed as a synonym).

Gnaphalium cylindrostachyum Wallich, Num.
List [Wallich] 2931 (1831), nomen nudum.

Illustrations : Cabrera & Ragonese (1978:
240); Porteners (1992: 204); Stanley & Ross
(1986: 532).

Weakly aromatic shrub 40-100 cm high; stems

Austrobaileya 8(3): 280-334 (2011)

one to several from non-woody rootstock,
sparsely to densely lanate and sometimes
with a few globose sessile glands; stem wings
entire, 1.4-3.3 mm wide. Leaves oblanceolate
to obovate, 16-45 x 6.5-16 mm, 2.1-3.4
times longer than wide, apex acute or obtuse;
margins recurved, denticulate, with 7-16
pairs of blunt teeth. Indumentum comprising
lanate multi-cellular hairs, glandular hairs
absent. Upper surface somewhat bullate;
hairs erect to spreading, sparse to moderately
dense, basal cells short; globose sessile
glands absent. Lower surface with lanate
hairs erect to spreading, moderately dense to
dense, extending 0.5-1 mm from the surface,
basal cells not very short; globose sessile
glands yellow and shining, dense, touching
or up to 3 diameters apart. Capitulescence
spiciform, 33-75 mm long and 8-14 mm wide
at anthesis, 2.8-5.8 times longer than wide;
side branchlets never continuous, at 20-45°;
peduncles 5-25 mm long. Outer involucral
bracts 1.6-2 mm long, oblong to oblong-
spathulate, with dense spreading lanate hairs
on outer surface, glabrous on inner surface;
glands absent; apex obtuse. Inner bracts white
to straw coloured (sometimes tinged pink) on
dorsal surface; longest ones linear, 3-3.6 x 0.1-
0.25 mm wide, apex attenuate, upper margins
entire or with an occasional small tooth; cilia
5-9, up to 2.5 mm long, attached along the
proximal half and consistently extending to
the distal one third, not exceeding the bract
apex. Filiform (female) florets 27-45, white
or straw-coloured, corolla 2.3-2.8 mm long,
styles extending 0.1-0.7 mm beyond corolla.
Hermaphrodite floret solitary, corolla pink,
2.5-3 mm long; corolla lobes 0.25-0.5 mm
long, globose sessile glands always present,
eglandular trichomes absent. Achenes
narrowly ellipsoidal, 0.5-0.7 mm long, terete
or slightly angular, dark brown, with 0-25 or
25-50 twin hairs each c. 0.05 mm long, evenly
distributed; globose sessile glands at junction
between achene and pappus usually present;
pappus 2.2-3 mm long. Figs. 11I-L, 13.

Additional selected specimens examined : China.
Ch’ang-kiang District, Hainan Island, Feb 1934, Lau
3261 (GH); Taichow, Hainan Island, Jan 1938, How &
Chun 70001 (GH). India. Ghariajhor, 11 miles [18 km]
from Sundargarh, Orissa, Mar 1950, Mooney 3748 (K).
Myanmar. Pegu, s.dat., Kurz 2263 (K); Prome, Ava,

Bean, Pterocaulon section Monenteles

307

QUEENSLAND HERBARIUM (BRI)

Flora of Queensland Maranoa

Pterocaulon reddens (Willd.) Fern.-Vill.

QUEENSLAND HERBARIUM (BRI)
Brisbane Australia

747G83

Fig. 13. Representative specimen of Pterocaulon redolens (Eddie CPE1064 [BRI]).

308

s.datHance 17401 (K). Thailand. Ban Huay Nam
Kow, Chiang Mai Province, Feb 1993, Radanachalesa
1 (A). Indonesia, near Palu, Celebes, Apr 1975, Meijer
9173 (A, L). Philippines. Bataan Province: Luzon, Jul
1913, Merrill 341 (AMES, L). Papua New Guinea.
Morobe Province: Erap, Jan 1958, Henty NGF9481
(BRI, CANB, K). Central Province: Tupuseleia, Port
Moresby subdistrict, Aug 1967, Kairo & Streimann
NGF30803 (BRI, CANB, L). New Caledonia. lie Pott,
Mouane, Aug 1968, McKee 19366 (NOU); Plateau
central, lie des Pins, Jul 1970, McKee 22304 (NOU);
Noumea, 1868-1870, Balansa (A). Vanuatu. Tableland,
Eromanga, Jul 1896, Morrison s.n. (K). Australia:
Queensland. Cook District: Dimbulah, Apr 1935,
Flecker 398 (CNS); Ravenshoe - Hughenden Road, c.
150 km N of Hughenden, Apr 1975, Halliday 418 (BRI);
Bulleringa N.P., 80 km NW of Mt Surprise, Apr 1998,
Forster PIF22541 & Booth (BRI, DNA, MEL). North
Kennedy District: Barrabas Scrub, May 1972, Hyland
6085 (CNS); 13.5 km W of the Dry Swamp, May 1991,
Neldner 3268 & Thompson (BRI, K); 6 miles [10 km] N of
Craigie Station, Jul 1953, Lazarides 3718 (BRI, CANB);
Townsville, Aug 1916, White s.n. (BRI [AQ270344]).
Leichhardt District: Springsure, Apr 1961, Jones 1870
(BRI, CANB); Twenty Mile Yards, ‘Rookwood’, Apr
1991, Forster PIF7881 & McDonald (BRI, MEL). Port
Curtis District: Rundle Range, S.F. 60, 37 km NW of
Gladstone, Aug 1989, Gibson TOI698 (BRI). Burnett
District: 9 km SE of Mt Perry (town), Apr 1985, Rodd
& Hardie 4378 (BRI, CANB, NSW); Windera, Apr
1963, McKee 10198 (BRI, CANB). Maranoa District:
Boatman Station, Mar 1947, Everist 2813 (BRI); ‘Six
Mile’, Bulloak Paddock, NNW of Roma, Mar 2007,
Eddie CPE1064 (BRI). Warrego District: Yanna
Siding, via Charleville, May 1949, Heinemann s.n. (BRI
[AQ270369]). Moreton District: Rifle Range Reserve,
Goodna, E of Ipswich, Apr 1990, Bean 1512 (BRI).

Distribution and habitat : Pterocaulon
redolens is widespread, with occurrences in
China (Hainan), India, Myanmar, Thailand,
Philippines, Indonesia, Papua New Guinea,
Australia, New Caledonia and Vanuatu.
Cabrera & Ragonese (1978) reported seeing
specimens also from Laos, Cambodia and
Vietnam. In Australia, it is widely distributed
in eastern Queensland, south from Lakefield
National Park (Map 6), and was reported by
Porteners (1992) to occur in the far north¬
east of New South Wales. It inhabits grassy
areas at low altitudes in hilly or flat terrain,
with sandy to clay-loam soil. In Australia, it
is found in various eucalypt woodlands or
on the edge of vine thickets in shallow or
deep soils, sometimes behind coastal dunes.
The geological substrate may be granite,
sandstone, basalt or serpentinite.

Austrobaileya 8(3): 280-334 (2011)

Phenology : Flowers and fruits are recorded
for every month of the year, with most records
between April and September.

Affinities: Pterocaulon redolens is not
obviously allied to any other species. However,
sterile specimens of Pterocaulon redolens
are indistinguishable from the sympatric
P. ciliosum.

Typification: The isotype of Conyza redolens
at K [K000373288] has a blue hand-written
label stating “Gnaphalium redolens (Forster)/
Habitat in New Caladonia”, and next to that, a
printed label saying “The Forster Herbarium.
Presented by the Corporation of Liverpool,
August 1885”. There are five separate branches
mounted on the sheet; three are Pterocaulon
redolens and two are P. sphacelatum.

Notes: This species is distinctive in the
spicate, interrupted capitulescence; ring of
glands often present at apex of achene; upper
margins of inner bracts entire and many
female florets (27-45) per capitulum.

The author has seen a single specimen
of this species from India, as cited above.
Pterocaulon redolens must be a relatively
recent arrival in India, as Hooker (1882)
recorded it only from “Burma and the Eastern
Peninsula”. It was not recorded by Haines
(1961) as present in Bihar or Orissa. Rath &
Priyadarshini (2005) reported Pterocaulon
redolens from an abandoned manganese
quarry, a highly modified habitat. I conclude
that this species is very likely not native to
India, but naturalised.

9. Pterocaulon serrulatum (Montrouz.)
Guillaumin, Bull. Soc. Bot. France 84: 56
(1937); Monenteles serrulatus Montrouz.,
Mem. Acad. Roy. Sci. Lyon , Sect. Sci. 10: 225
(1860). Type: New Caledonia. Art Island,
s.dat., J.X.Montrouzier 125 (holo: LY,
destroyed).

Monenteles glandulosus Benth., FI. Austral.
3: 523 (1867); Pterocaulon glandulosum
(Benth.) F.Muell, Syst. Census Austral. PI.
79 (1883); P. glandulosum var. glandulosum ,
Ewart & Davies, FI. N. Territory 277 (1917).
Type: Australia: Queensland. Cook District:
Gilbert River, October 1856, F. Mueller s.n.
(lecto: K373279, fide Cabrera & Ragonese
[1978: 254]).

Bean, Pterocaulon section Monenteles

309

Strongly aromatic perennial shrub 35-150 cm
high; stems oneto several from woody rootstock,
with sparse to dense lanate eglandular hairs,
globose sessile glands and sometimes stalked
glandular hairs; stem wings serrate, 2.5-8
mm wide. Leaves elliptical, ovate or broadly-
lanceolate, 20-82 x 8-28 mm, 2.1-4.3 times
longer than wide, apex acute; margins flat or
recurved , serrulate, with 13-45 pairs of teeth.
Indumentum comprising lanate multi-cellular
hairs, glandular hairs present or absent. Upper
surface smooth or slightly bullate; hairs erect
to spreading, sparse to dense, basal cells
short; globose sessile glands present. Lower
surface with lanate hairs erect to spreading,
sparse to very dense, extending 1-1.5 mm
from the surface, basal cells not very short;
globose sessile glands yellow and shining or
pale yellow and opaque, dense, touching or up
to 3 diameters apart. Capitulescence ellipsoid
or cylindrical, sometimes ‘interrupted’, rarely
almost globose, 14-52 mm long and 13-17
mm wide at anthesis, 1.1-3.5 times longer
than wide; side branchlets never continuous,
at 20-45°; peduncles 0-15 mm long. Outer
involucral bracts 2.3-37 mm long, oblong,
with sparse to moderately dense spreading
lanate hairs on margins, glandular-hairy on
both surfaces; apex obtuse. Inner bracts white
to greenish-white on dorsal surface; longest
ones linear, 3.9-6.3 x 0.3-0.5 mm, apex acute,
upper margins with small or large teeth; cilia
6-14, up to 2.4 mm long, attached along the
proximal half and consistently extending to
the distal one third, not exceeding the bract
apex. Filiform (female) florets 29-52, white
to straw coloured, corolla 2.8-4.2 mm long,
styles extending 0.3-0.5 mm beyond corolla.
Hermaphrodite floret solitary, corolla pink,
3.6-47 mm long; corolla lobes 0.4-0.6
mm long, globose sessile glands present,
eglandular trichomes absent. Achenes
narrowly ellipsoidal, 0.7-1 mm long, terete
or almost so, dark brown, with 25-50 twin
hairs each c. 0.1 mm long, evenly distributed;
globose sessile glands at junction between
achene and pappus absent; pappus 2.8-4.5
mm long.

Affinities : Pterocaulon serrulatum is

a distinctive species, but is sometimes
confusable with P. sphacelatum and
P. ciliosum. These are perhaps its closest

relatives, and postulated hybrid specimens
are reported in this paper.

Typification: Vegter (1976) reported that
Montrouzier’s types are at LY, and Guillaumin
& Beauvisage (1914) reported seeing the type
of Monenteles serrulatus there. However, it
appears that Montrouzier’s collection at LY
was destroyed or lost in the 1950’s (P. Morat,
pers. comm. June 2010), and the curator at LY
has confirmed that the type of Monenteles
serrulatus is not present there now. Duplicates
of some of Montrouzier’s collections are held
at MPU, but no type material of Monenteles
serrulatus can be found there (P. Schafer,
pers. comm. June 2010), nor is there an isotype
at P (P. Morat, pers. comm. June 2010). This
is in accord with the information provided
by Guillaumin & Beauvisage (1914), i.e. that
there was only a single type specimen for this
name, the specimen at LY.

There seems little doubt about the
application of the name Pterocaulon
serrulatum. The protologue is fairly detailed
and the description matches the plant currently
called Pterocaulon serrulatum var. serrulatum
in Australia. Guillaumin & Beauvisage (1914)
reported that the type was very similar to
Pterocaulon glandulosum F.Muell., but that it
differed by being totally glabrous.

Notes : Pterocaulon serrulatum is distinctive
in the cylindrical to ovoid capitulescence;
broad serrated stem wings; many female
florets per capitulum; outer involucral bracts
glandular hairy on both surfaces and the
sessile glands on upper leaf surface.

Varietal rank is appropriate for the taxa
included within Pterocaulon serrulatum. The
two varieties differ only in their indumentum,
but these characters do not vary continuously
in Australian material, and a useful
geographical separation is achieved through
their use. All eastern Queensland material can
be readily assigned to var. serrulatum , and all
Northern Territory material is referrable to a
more broadly circumscribed var. velutinum,
as outlined in the key. Some specimens from
the Kimberley region are assigned here to var.
serrulatum. Some specimens from the Jericho
and Hughenden districts of Queensland are
difficultto assign to either variety; for example,

310

Austrobaileya 8(3): 280-334 (2011)

QUEENSLAND HERBARIUM (BRI)

Flora of Queensland Leichhardt

Pterocaulon serrulatum (Montrouz.) Guillaumin
var. serrulatum

QUEENSLAND HERBARIUM (BRI)
Brisbane Australia

aq 649945

(55,512865,7294521 ) (8349-128945) Alt. 400m

118km from Spnngsure, on road to Tambo.

Low woodland or shrubland with Acacia harpophylla, Erythroxylum.
Eremophila mitchellii. Carissa Pink loam near crest of low ridge.
Highly aromatic forto, stems and leaves green, Inflorescence mauve.
Rare at site

Fig. 14. Representative specimen of Pterocaulon serrulatum var. serrulatum ( Bean 22120 [BRI]).

Bean, Pterocaulon section Monenteles

311

Smyrell GS69 has broad based glandular
hairs (a feature of var. serrulatum) and also
has lanate hairs on the upper leaf surface (a
feature of var. velutinum). Similarly, in the

Pilbara region of Western Australia, there are
specimens referred to here as var. velutinum ,
but which closely approach var. serrulatum
e.g. Byrne 2807.

Key to varieties

Lanate hairs absent from upper leaf surface; broad based glandular hairs

common on leaves, stems and stem wings.

.9a. P. serrulatum var. serrulatum

Lanate hairs present on upper leaf surface; broad based glandular trichomes
absent, but globose sessile hairs common on leaves, stems and stem wings
.9b. P. serrulatum var. velutinum

9a. Pterocaulon serrulatum var.
serrulatum

Winged stems and leaf surfaces with many
broad-based glandular hairs 0.1-0.2 mm long;
erect or spreading eglandular multicellular
lanate absent from upper leaf surface, sparse
or absent on lower leaf surface, stem wings
and stems. Figs. IB, 14,16A-D.

Illustration : Stanley & Ross (1986: 532).

Additional selected specimens examined : Australia:
Western Australia. Bell Gorge parking area, Aug
2005, Byrne 1619 (PERTH); western base of Mt
Hamilton, Jul 1905, Fitzgerald 1209 (PERTH); Little
Island, off Long Island, Buccaneer Archipelago, Jun
1982, Hopkins BA0115 (PERTH); Around ‘Beverley
Springs’ Homestead buildings, Jun 1992, Mitchell 2556
(PERTH). Queensland. Cook District: Fossilbrook
Creek, ‘Burlington’, N of Mount Surprise, Aug 1997,
Bean 12229 (BRI); along Kuranda Highway, c. 10
miles [16 km] E of Mareeba, Jun 1962, Hoogland 8486
(BRI, CANB). North Kennedy District: 22.5 km (by
road) SW of Herberton, on the Silver Valley road.
May 1983, Conn & Clarkson 1178 (AD, BRI, CANB,
CNS, DNA, HO, NSW, PERTH); Road from Camel
Creek to Greenvale, Aug 1998, Fox 25 & Bean (BRI);
base of Frederick Peak, 25 km SW of Townsville, May
1991, Bean 3215 (BRI); Lookout, on eastern slopes of
Mt Kelly, c. 14 km south-west of Ayr, May 2009, Bean
28848 (BRI); Bowen, reserve near reservoir, Nov
1978, Ross s.n. (BRI [AQ297444]). Mitchell District:
‘Grant’, c. 70 km ENE of Barcaldine, Jul 1994, Smyrell
GS89 (BRI). South Kennedy District: Redcliff Island,
Seaforth, Aug 1992, Batianoff9208129 (BRI); 19 km E
of‘Doongmabulla’, N of Jericho, Apr 2002, Bean 18984
(BRI, MEL). Leichhardt District: 60 miles [97 km] N
of Comet township, Jul 1962, Story & Yapp 185 (BRI,
CANB); 118 km from Springsure, on road to Tambo,
May 2004, Bean 22120 (BRI); Ka Ka Mundi section,
Carnarvon N.P, on fire trail from Salvator Rosa, Aug
2008, Thomas MBT3780 & Leggett (BRI). Port Curtis
District: ‘Corisande Hills’, Stanage Bay road, N of
Rockhampton, Jul 1997, Bean 12063 (BRI, MEL); 32

km N of Miriam Vale, Jul 1996, Thompson M1R136A
& Turpin (BRI). Burnett District: Gayndah, May
1917, White s.n. (BRI [AQ2703920]); ‘Dykehead’, 70
km W of Mundubbera, Jun 1998, Robinson JR-1 (BRI).
Wide Bay District: Degilbo Creek, near Biggenden,
Aug 1980, Johnson s.n. (BRI [AQ343756]); Rossmore
Road, 10 km SE of Kilkivan, May 2006, Webb s.n. (BRI
[AQ619054]).

Distribution and habitat: Found on Art
Island, New Caledonia, and throughout the
eastern half of Queensland in Australia,
from Cooktown to Kilkivan. It also occurs
in the western Kimberley region of Western
Australia (Map 7). It grows on a wide range
of sites (include coastal headlands) where
drainage is good, usually in eucalypt woodland
or grassland. Soils range from sands to clay-
loams.

Phenology: Flowers and fruits are borne
between April and November.

9b. Pterocaulon serrulatum var. velutinum
(Ewart & O.B.Davies) Guillaumin, Bull.
Soc. Bot. France 84: 57 (1937); Monenteles
glandulosum var. velutinum Ewart &

O. B.Davies, FI. N. Territory 211 (1917). Type:
Australia: Northern Territory. Haast’s Bluff,
17 May 1911, G.F.Hill 184 (holo: MEL).

P. serratum O. Schwarz, Repert. Spec. Nov.
Regni Veg. 24: 108 (1927). Type: Australia:
Northern Territory. Port Darwin, 10 miles
NE, s.dat., F.A.K.Bleeser 712 (holo: B,
destroyed).

Illustrations: Cabrera & Ragonese (1978:
253); Wheaton (1994: 112); Barrs (1999: 118)
as P. serrulatum var. serrulatum.

312

Austrobaileya 8(3): 280-334 (2011)

QUEENSLAND HERBARIUM <BRI)

Flora of Queensland Burke

Pterocaulon serrulatum (Montrouz.) GuiNaumin

Coil. R. Booth LH28-5 7 JUL 2009

Kelman.0.:

(10d 57m 50s 139d 3m 08s) [WGS84J Depth m

54.294910.7902039 (6760-94920) AIL m

Gregory Downs Stn, E of Rrversleigh.

Eucalyptus leucophfoia woodland on Jurrasic sandstone ridgetop.
Hairy upright herb.

Queensland Herbarium (BRI)

Pterocaulon serrulatum var uelutin,

CE-tiaanwajSSSr

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

850132

Fig. 15. Representative specimen of Pterocaulon serrulatum var. velutinum {Booth LH28-5 & Kelman [BRI]).

Bean, Pterocaulon section Monenteles

313

Fig. 16. Pterocaulon serrulatum var. serrulatum. A. hermaphrodite floret x 32. B. inner involucral bract (cilia not
shown) x 16. C. upper portion of inner involucral bract x 32. D. mature achene and basal portion of pappus x 32.
Pterocaulon sphacelatum. E. hermaphrodite floret x 32. F. inner involucral bract (cilia not shown) x 16. G. upper portion
of inner involucral bract x 32. H. mature achene and basal portion of pappus x 32. Pterocaulon sphaeranthoides. I.
hermaphrodite floret x 32. J. inner involucral bract (cilia not shown) x 16. K. upper portion of inner involucral bract
x 32. L. mature achene and basal portion of pappus x 32. A-C from Bean 28848 (BRI); D from Ross s.n. (BRI
[AQ297444]); E, H from Piercey 51 (DNA); F-G from Halford Q2626 (BRI); I-K from van Leeuwen 0099 (PERTH);
L from van Leeuwen 0092 (PERTH).

314

Winged stems and leaf surfaces without
broad-based glandular hairs; globose sessile
glands c. 0.05 mm long, abundant; erect to
tangled eglandular multicellular lanate hairs
sparse to very dense on leaves, wings and
stems. Figs. 1A, 15.

Additional selected specimens examined: New
Caledonia. Voh, presqu’ile de Gatope, Aug 1970,
McKee 22393 (NOU); Mt Yallein, Jul 1968, Jaffre
57 (NOU); Presqu’ile d’Arama, Aug 1965, Schmid
619 (NOU); West face of Massif de Koniambo, Oct
1963, Green 1288 (A, K, NOU). Australia: Western
Australia. Gibb River Road, 117 km E of Derby, Aug
2001, Courtney 40 (PERTH); Honeymoon Beach, 25
km N of Kalumburu, Jun 1990, Edinger 713 (PERTH);
37.6 km NW of ‘Bonney Downs’ Homestead on road to
Hillside, Oct 1995, Mitchell PRP971 (DNA, PERTH);
McLarty Hills, Great Sandy Desert, Aug 1977, George
14625 (CANB, PERTH); 12 km SSW of Two Sisters,
c. 150 km SE of Shay Gap, Jul 1984, Newbey 10407
(PERTH); E of junction of Kununurra - Wyndham
-Halls Creek roads, Jul 1974, Carr 3203 & Beauglehole
46981 (BRI, CANB, PERTH); Roy Hill, Aug 2007, Byrne
2807 (PERTH); Meentheena Conservation Reserve, 6.5
km SSE of ‘Meentheena’ Homestead, May 2001, van
Leeuwen 4797 (PERTH). Northern Territory. 4.5 km
from Stuart Highway on Edith Falls Road, Jul 2003,
Elick 222 (CANB, CNS); 23 miles [37 km] NW of Rabbit
Flat, Tanami, Apr 1971, Dunlop 2131 (BRI, DNA); base
of Meyers Hill, Alice Springs, Aug 1957, Chippendale
3587 (BRI, CANB, DNA); Mt Bundy, Oct 1968, Harpley
NB982 (BRI, DNA); 269 km N of ‘Barkly’ Homestead
towards Borroloola, Jun 1999, Bean 15047 (BRI, DNA);
Channel Island, Darwin Harbour, Jul 1995, Cowie 5880
(DNA, MEL). Queensland. Burke District: 3 miles [5
km] from Mount Isa towards Lake Moondarra, Jul 1974,
Ollerenshaw 1142 & Kratzing (BRI, CANB, L); Gregory
Downs Station, E of Riversleigh, Jul 2009, Booth LH28-

5 & Kelman (BRI). Cook District: Ortona Station,
102 km by road S of Forsayth, Aug 2010, Bean 29878

6 McDonald (BRI, MEL, NSW). Mitchell District:
Bessie’s Castle, Silsoe Road, c. 100 km W of Longreach,
May 2004, Bean 22246 (BRI); 19.7 km E of Jericho, Jul
1975, Chapman 1281 (AD, BRI, CANB, K, L). Gregory
North District: Winton - Jundah Road, 23.9 km N of
‘Elvo’ Homestead, May 2004, Bean 22561 (BRI).

Distribution and habitat : A widespread
taxon, occurring in New Caledonia, Australia
in central and north-western Queensland,
throughout the Northern Territory, and the
northern half of Western Australia as far
west as the Hamersley Range (Map 7). It is
also recorded for a few locations in northern
South Australia (Anon. 2011). It inhabits a
wide range of habitats; the vegetation can be
Acacia woodland, mallee shrubland, eucalypt
open woodland or Triodia grassland. Soils can
vary greatly, including beach sand, red sandy

Austrobaileya 8(3): 280-334 (2011)

loam, or basaltic clay-loams, and it may grow
on sandstone, limestone or quartzite hills, or
on flats.

Phenology : Flowers and fruits have been
recorded from April to October.

Notes : Cabrera & Ragonese (1978) stated
that Pterocaulon glandulosum var. velutinum
Ewart & O.B.Davies is a nomen nudum , but
that is incorrect. Ewart and Davies provided
an adequate description and the name is
validly published.

The type of Pterocaulon serrulatum var.
velutinum has a very dense cover of lanate
eglandular hairs, and the two varieties have
in the past been distinguished on the density
of these hairs (based on herbarium specimen
determinations). In this paper, I have placed
more emphasis on the presence or absence
of stalked glandular hairs in dividing the
varieties. This provides a useful geographic
separation between the varieties, but it means
that a wide range of lanate hair densities is
included within var. velutinum. The density of
lanate hairs can vary at a single location, e.g.
Latz 7950 comprises pieces collected from
two plants growing at the same place; on one
the lanate hairs are sparse, and on the other
they are very dense.

No type material of Pterocaulon serratum
O. Schwarz has been seen, but from the
description given in the protologue, there
seems little doubt that it is this variety.

10. Pterocaulon sphacelatum (Labill.)
F.Muell., Syst. Census Austral. PI. 79 (1882);
Monenteles sphacelatus Labill., Serf.
Austro-Caledon. 43 (1825). Type: [New
Caledonia] Nova Caledonia. [June 1792],
J.J.H.Labillardiere s.n. (lecto: FI [FI-W
92608], here designated, image!; isolecto:
FI [FI-W 92605, left hand specimen only],
P537804, image!).

Illustrations : Cabrera & Ragonese (1978:
246); Porteners (1992: 204); Milson (2000:
21); Moore (2005: 126).

Strongly aromatic shrub 40-100 cm high,
often almost as wide; stems single or
numerous, densely lanate and sometimes
with a few globose sessile glands; stem wings
entire or sparsely toothed, 0.7-2 mm wide.

Bean, Pterocaulon section Monenteles

315

Leaves oblanceolate or spathulate, 20-66 x
4-15 mm, 3.3-5.8 times longer than wide,
apex obtuse or acute; margins recurved,
denticulate, with 6-14 pairs of blunt teeth.
Indumentum comprising lanate multi-cellular
hairs, glandular hairs absent. Upper surface
moderately to strikingly bullate; hairs erect
to spreading, sparse to moderately dense,
basal cells short; globose sessile glands
absent. Lower surface with lanate hairs erect
to spreading, moderate to dense, extending
0.4-0.7 mm from the surface, basal cells not
very short; globose sessile glands yellow and
shining, dense, touching or up to 3 diameters
apart. Capitulescence globose or ellipsoidal,
10-15 mm long, 10-14 mm wide at anthesis,
1-1.4 times longer than wide; side branchlets
never continuous, at 30-60°; peduncles 0-22
mm long. Outer involucral bracts 2.2-37 mm
long, spathulate, with dense spreading lanate
hairs throughout on the outer surface, sparse
hairs at distal end of the inner surface; glands
absent; apex acute. Inner bracts violet, pink or
white on dorsal surface; longest ones linear,
3.5-4.6 x 0.3-0.5 mm; apex acute, upper
margins conspicuously toothed, or upper
margins lacerate with no single distinct apex;
cilia 6-18, up to 1.7 mm long, mostly attached
in proximal half and few extending to the distal
one third. Filiform (female) florets 17-29, pink,
corolla 2.4-3.2 mm long, styles extending
0.4-0.6 mm beyond corolla. Hermaphrodite
floret solitary, corolla pink, 2.5-37 mm long;
corolla lobes 0.4-0.6 mm long, globose sessile
glands present, eglandular hairs absent.
Achenes narrowly ellipsoid, 0.7-1 mm long,
terete or slightly quadrangular, dark brown,
with 25-50 twin hairs each c. 0.05 mm long,
evenly distributed; globose sessile glands at
junction between achene and pappus rarely
present; pappus 2.3-3.3 mm long. Fruit salad
plant, Apple bush. Figs. 2A, 16E-H, 17.

Additional selected specimens examined : Timor-
Leste. Plateau of Baucau, Dec 1953, Van Steenis 18030a
(L). New Caledonia. Baie Maa, Oct 1982, Suprin
2059 (NOU); s.loc. , Oct 1965, MacKee 13525 (K, L).
Australia: Western Australia. Eagle Bore Study site.
Plot 9, Gibson Desert Nature Reserve, Aug 2002, Bragg
118 (MEL, PERTH); Rocky Pool, Gascoyne River, c.
850 km N of Perth, Oct 1975, Kenneally 4641 (PERTH);
Northern end of Windjana Gorge, Jul 1974, Carr 4281
& Beauglehole 48059 (PERTH); Elder Creek, 2 km
W of Warburton, Aug 1962, George 3828 (PERTH);
Anna Plains Station, Jul 1941, Burbidge 1462 (PERTH);

Lacrosse Island, at head of Cambridge Gulf, Jun 1992,
Kenneally 11328 (DNA, CANB, PERTH). Northern
Territory. Auvergne Station, 35 km NW of Timber
Creek, Jul 1977, Must 1620 (CANB, DNA, SI); c. 4.5 km
by road N of Gilbert Creek crossing by Stuart Highway,
between Wauchope and Tennant Creek, Aug 1978,
Donner 6232 (AD, CANB, NT); Junction Reserve, Jul
1982, Piercey 51 (DNA); Oenpelli, Sep 1948, Specht
1083 (BRI, L); Burt Creek, 4 miles [6 km] S of Yambah
Station, Mar 1953, Perry 3372 (BRI, CANB); 3.2 miles
[5.1 km] N of Alice Springs, Dec 1962, Nelson 559
(CANB, DNA); Andado Station, Mac Clarke Reserve,
Sep 1992, Latz 12706 (DNA, MEL); Horse paddock,
‘Tobermorey’, Oct 1954, Chippendale 418 (DNA); Barkly
Tablelands, N of Telegraph bore, Mittibah Station, Jul
2001, Mangion 1534 & Risler (DNA). Queensland.
Burke District: East bank of Leichhardt River, 72 km
SE of Burketown, Burke Development Road, Jul 1991,
Jobson 1490 (BRI, HO, MEL). Cook District: Topsy
Creek, Kowanyama Aboriginal Reserve, Aug 1980,
Clarkson 3392 (BRI, CNS). Gregory North District:
32 km N of Bedourie along road to Boulia, Jul 1995,
Halford Q2626 (BRI). Mitchell District: Silsoe Road,
4.1 km W of ‘Morelia’ turnoff, W of Longreach, May
2004, Bean 22250 (BRI). Warrego District: 99.3 km
from Blackall towards Adavale, near Listowel Downs
Station, Oct 1983, Canning 6192 & Rimes (CANB, BRI,
DNA, MEL, NSW). Gregory South District: 20.2 km
WSW of Eromanga, on Cooper Developmental Road,
Aug 2010, Bean 30023 (A, BRI, BRIT, MO). New South
Wales. 113 km W of Wanaaring on Milparinka Road,
Nov 1971, Blaxell 613 (BRI, NSW); Lake Pinaroo near
Fort Grey camping area, NW of Tibooburra, Sep 1989,
Coveny 13477 et al. (AD, BRI, CANB, NSW, MEL,
PERTH); Homestead Gorge, Mootwingee N.P, 112
km NE of Broken Hill, Oct 1988, Crawford 1028 (AD,
CANB, NSW). South Australia, around Nent Oura
Research unit. Mount Freeling Station, Oct 1987, Foster
227 (AD, DNA); Stuart Creek, 18 km WSW of ‘Stuart
Creek’ Homestead, Dec 1984, Badman 1597 (AD, DNA);
Bunyeroo Gorge, Flinders Ranges N.P, Nov 1982,
Symon 13040 (AD, CANB, MO).

Distribution and habitat : Pterocaulon
sphacelatum is the most widespread
Australian Pterocaulon species, occurring
in all mainland states except Victoria, and
extending from the west coast of Western
Australia to within 150 km of the Queensland
east coast. It occurs throughout much of
Northern Territory and South Australia, and
in the north-west of New South Wales. It also
occurs in Timor and New Caledonia (Map
8). In grows on a range of sandy to clay-
loam soils, on stony hillsides (higher rainfall
areas) or creek-beds (arid areas), in grassland
(often dominated by Triodia spp.), eucalypt
woodland or low open woodland with Acacia
aneura or other Acacia spp.

316

Austrobaileya 8(3): 280-334 (2011)

oriA

W* m.

northern territory herbarium

DARWIN (DIVA) AUSTRALIA
Pterocsmlon sphacelatum (Labill.) Benlh. & Hook, si F.Muell

Barkly Tablelands. N.of Telegraph bore Millibah Slalion

STATE/DIST.: NT /BT I8"4479"S 137=I710"E

Perennial herb lo 50cm tall on cresl of gravelly clay soil on plain.

Fig. 17. Representative specimen of Pterocaulon sphacelatum (Mangion & Risler 1534 [DNA]).

Bean, Pterocaulon section Monenteles

317

Phenology : Flowers and fruits occur from
June to October in northern parts of its range,
and from August to December in southern
parts.

Affinities : Pterocaulon sphacelatum is
closely related to P. ciliosum, but differs by
the narrower leaves, and the bright pink inner
bracts with few visible cilia on protruding
parts, and strongly toothed to lacerate upper
margins.

Typification: There are two sheets at FI
bearing material that matches the protologue
of Monenteles sphacelatus. On one sheet (FI-
W 92605), specimens of both Pterocaulon
sphacelatum and P. redolens are present, but
neither specimen is named; on the other sheet
(FI-W 92608), there is a single specimen of
P. sphacelatum accompanied by a detailed
description written by Labillardiereandheaded
Monenteles sphacelatus ; this description
agrees perfectly with the protologue. This
latter sheet is selected as the lectotype.

Notes : This species is distinctive in the upper
margins of the inner bracts often lacerate;
capitulescences often ellipsoidal, up to 1.4
times longer than wide; the inner bracts always
partly pink to violet, often predominantly so;
leaves oblanceolate in shape, 3.5-5.8 times
longer than wide; leaf upper surface bullate;
corolla lobes of hermaphrodite floret short
and none or few cilia visible on protruding
part of inner bracts.

11. Pterocaulon sphaeranthoides (DC.)
F.Muell., Syst. Census Austral. PI. 79 (1882);
Monenteles sphaeranthoides DC., Prodr. 5:
456 (1836). Type: Australia: Western Australia.
Enderby Island, Dampier Archipelago, 25
February 1818, A.Cunningham s.n. (holo: G-
DC; iso: PERTH; BM, K n.v).

Illustration : Cabrera & Ragonese (1978:
250).

Strongly aromatic shrub 30-120 cm high;
stems numerous, densely glandular-hairy
and sometimes with sparse lanate eglandular
hairs; stem wings prominently toothed,
0.5-3(-4) mm wide. Leaves narrowly
oblanceolate to linear, 12-48 x 2-10 mm,
3.5-13 times longer than wide, apex acute;
margins recurved to revolute, serrate with

10- 16 pairs of teeth. Indumentum comprising
broad-based glandular hairs, globose sessile
glands and sometimes lanate eglandular hairs.
Upper surface strikingly bullate; glandular
hairs erect, sparse to moderately dense,
0.1-0.2(-0.4) mm long, broad-based; lanate
hairs absent or sparse; globose sessile glands
present. Lower surface glabrous or with
sparse, erect to spreading lanate hairs, basal
cells not very short; globose sessile glands
yellow and shining, dense, touching or up to
3 diameters apart. Capitulescence ± globose,

11- 16 mm long, 11—14 mm wide at anthesis,
1-1.2 times longer than wide; side branchlets
never continuous, at 30-60°; peduncles 1-20
mm long. Outer involucral bracts 2.2-27
mm long, linear-spathulate to oblong, with
dense spreading lanate hairs throughout on
the outer surface, glabrous on inner surface;
glands present; apex acute or obtuse. Inner
bracts violet, pink or white on dorsal surface;
longest ones linear, 3.5-4 x 0.2-0.5 mm,
upper margins with large teeth, or lacerate
with no distinct single apex; cilia 5-12, up
to 1.5 mm long, mostly attached in proximal
half and few extending to the distal one third.
Filiform (female) florets 14-20, pink, corolla
2.3-2.8 mm long, styles extending 0.2-0.9 mm
beyond corolla. Hermaphrodite floret solitary
or very rarely two, corolla pink, 27-3.4 mm
long; corolla lobes 0.4-0.5 mm long, globose
sessile glands present, eglandular trichomes
absent. Achenes narrowly ellipsoid, 0.8-
0.95 mm long, slightly quadrangular, dark
brown, with 25-50 twin hairs each 0.05-0.1
mm long, evenly distributed; globose sessile
glands at junction between achene and pappus
occasionally present; pappus 2.2-2.9 mm
long. Figs. 16I-L, 18.

Additional specimens examined : Australia: Western
Australia. Hermite Island, Montebello Islands, Oct
2000, Kenneally 11558 (PERTH); Depuch Island, NE of
Roebourne, May 1962, Royce 7131 (PERTH); Charles
Knife Road, between picnic area and Cape Range N.P,
Aug 1978, Perry 845 (BRI, PERTH); Legendre Island,
Dampier Archipelago, Jun 1962, Royce 7264 (PERTH);
Barrow Island, Aug 1973, Butler 47 (PERTH); Rosemary
Island, Dampier Archipelago, Jun 1962, Royce 7451
(PERTH); Barrow Island Nature Reserve, 11.3 km
WNW of Town Point, Nov 1991, van Leeuwen 1086
(PERTH); 48.6 km NW of ‘Bonney Downs’ Homestead
on road to Hillside, Nov 1995, Mitchell PRP977 (DNA,
PERTH); Lower end of Upper Bee Gorge, Hamersley
Range N.P, Aug 1973, Weston 8443 (PERTH); Python

318

Austrobaileya 8(3): 280-334 (2011)

WESTERN AUSTRALIAN HERBARIUM, PERTH
Flora of Western Australia

Pterocaulon sp ha era ti th oides (DC.) F.Muell.

Astcraceae
Erect perennial herb.

Rock and proximal colluvium. Steep slope, aspect 330 degrees.
Many coarse fragments to maximum size of 200 mm. Abundant
bedrock outcrop. Brown loam, average depth 12.44 cm.
Scattered Low Trees of Terminalia canescens over Low
Scattered Shrubs over Hummock Grassland of Triodia sp. over
Very Open Herbs of Ptilotus exaltatus.

Coil. S. van Leeuwen et al. 0098 Date: 21/08/2005

Voucher: Pilbara Biological Survey

Loc.: Site: DRW05, West side of Karratha Stn HMS access
track, l.S km S of North West Coastal Highway. 9.2 km SW of
Mt Regal, 19.9 km SSW of Dampicr, Karratha Station, Pilbara
1BRA WA

Lai. 2G°51'14.300 W S Long. 116°40'6.700"E GDA94

Fig. 18. Representative specimen of Pterocaulon sphaeranthoides (van Leeuwen et al. 0098 [PERTH]).

Bean, Pterocaulon section Monenteles

319

Pool, Chichester-MillstreamN.P., Oct 1989, Nordenstam
& Anderberg 302 (PERTH); Yardie Creek, Cape Range
N.P., Dec 1986, Alford 816 (PERTH); Cleaverville Creek,
Roebourne, Jul 2004, Byrne 1075 (PERTH); W of Mt
Leal, on Roebourne to Wittenoon Highway, Sep 1981,
Lander 1132 (PERTH); Karrawingina Pool, Harding
River, Sep 1981, Weston 12772 (PERTH); Site OYE04,
3.4 km E of Pannawonica - Cape Lambert railway
crossing on Pannawonica - Midstream Road, Aug 2006,
van Leeuwen et al. 0102 (PERTH); Site PW17, 13.4 km
N of ‘Mt Flora’, 71.1 km E of Pannawonica, Sep 2006,
van Leeuwen et al. 0092 (BRI, PERTH); Durba Hills,
Heartland district, Jun 1984, Morse 180 (CANB); 3.1 km
SSE of Pulgorah Cone, Warrawagine Station, Pilbara,
Aug 2004, van Leeuwen 0090 (PERTH); West side of
Karratha Station Homestead access track, 1.8 km S of
North West Coastal Highway, Aug 2005, van Leeuwen
0098 (PERTH); North side of Fortescue River Mouth
access track, 6.8 km W of North West Coastal Highway,
Aug 2005, van Leeuwen 0099 (PERTH).

Distribution and habitat : This species is
endemic to Western Australia. It is common
along coastal parts of the Pilbara region,
including offshore islands, from the Cape
Range to Depuch Island, east of Roebourne.
It also occurs away from the coast, to the
south and south-east of Port Hedland, as far
east as the Durba Hills (Map 3). It grows in
Triodia grassland, or in Acacia shrubland
with Triodia understorey, on coastal sand
dunes, rocky hillsides or rocky watercourses.
Soils are often red sandy-loams. The substrate
may be limestone, sandstone or granite.

Phenology : Flowers and fruits have been
recorded from June to December.

Affinities : This species has affinities with
Pterocaulon sphacelatum, however, that
species has stem wings that are entire or
sparsely toothed, lanate hairs frequent and
glandular hairs absent from the upper leaf
surface.

Notes: This species is distinctive in the
blistered bright green narrow leaves; serrated
stem wings; lanate hairs absent or sparse and
the short broad-based glandular hairs present
on upper leaf surfaces.

Pterocaulon sphaeranthoides and
P. sphacelatum are very closely related
and some specimens are hard to identify,
suggesting that they do intergrade in some
areas. For example, Trudgen & Parnell 11608
(PERTH), Backhouse et al. BEM50 (PERTH),
and Lander 1132 (PERTH), all from the

Hamersley Ranges, while assigned here to
Pterocaulon sphaeranthoides , are also close
to P. sphacelatum. In other areas, they are
apparently sympatric; typical specimens of
both species have been seen from Barrow
Island, for example.

I have been unable to confirm the
observation made first by Bentham
(1867) that the involucres of Pterocaulon
sphaeranthoides have “Usually 2, very rarely
1 or 3” disc (hermaphrodite) florets. Cabrera
& Ragonese (1978) stated “male florets 1-
3”, as did Wilson (1992), both presumably
paraphrasing Bentham. Of the hundreds of
capitula examined by the present author,
all except one had a solitary hermaphrodite
floret. The exceptional capitulum had two
hermaphrodite florets.

12. Pterocaulon tricholobum A.R.Bean
species nova affinis P. globulifloro sed alis
caulinis multo latioribus, bracteis interioribus
longioribus, pilis numerosis in lobis
corollarum flosculorum hermaphroditorum,
achenarum pilis paucioribus et pappis
longioribus differens. Typus: Australia:
Northern Territory. East Alligator, 10 June
1971, G.C.Taylor 75 (holo: BRI; iso: CANB,
DNA).

Aromatic shrub 50-120 cm high; stems one to
several from woody rootstock, densely lanate
but without globose sessile glands; stem wings
entire, 2.8-5.5 mm wide. Leaves elliptical to
obovate, 25-92 x 8-30 mm, 2.1-3.7 times
longer than wide, apex obtuse or rarely
acute; margins recurved, denticulate, with
15-30 pairs of very small teeth. Indumentum
comprising lanate multi-cellular hairs,
glandular hairs absent. Upper surface not
or slightly bullate; hairs erect to spreading,
sparse to moderately dense, basal cell short;
globose sessile glands absent. Lower surface
with lanate hairs erect to spreading, dense to
very dense, extending 0.4-0.8 mm from the
surface, basal cells not very short; globose
sessile glands yellow and shining, moderately
dense, 3-6 diameters apart. Capitulescence
globose to broadly ellipsoidal, rarely
truncate-globose, (8—)10—16 mm long and 10-
14 mm wide at anthesis, (0.7-)0.9-1.3 times
longer than wide; side branchlets very often

320

Austrobaileya 8(3): 280-334 (2011)

270504

tiObOTyof Q ueenslancl Herbarium (BRI)

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HERBARIUM OF NORTHERN TERRITORY

Alice Springs, Australia
N.T.33145.

Pterooaulon sphaeranthoides (DC.) F. Muell,

Loc. Bast Alligator.

Coll. G * c * Taylor 75 Dote 10 June 1971

Notes Erect plant to 36cms. Growing

amongst rooks in sandy grey soil.

Fig. 19. Holotype of Pterocaulon tricholobum ( Taylor 75 [BRI])

Bean, Pterocaulon section Monenteles

321

continuous, at 0-80°; peduncles often absent,
but sometimes up to 11 mm long. Outer
involucral bracts oblong-spathulate, 2.5-3
mm long, with dense spreading lanate hairs
on outer surface, sparse to moderately dense
hairs on the distal end of inner surface; glands
absent; apex obtuse. Inner bracts white, rarely
tinged pink on dorsal surface; longest ones
linear, 4-4.8 x 0.2-0.35 mm, apex acute,
upper margins with some small or large teeth;
cilia 11-22, up to 2.5 mm long, attached along
the proximal half and consistently extending
to the distal one third, not exceeding the bract
apex. Filiform (female) florets 15-21, white
or straw-coloured, corolla 2.3-2.9 mm long,
styles extending 0.6-1 mm beyond corolla.
Hermaphrodite floret solitary, corolla pink,
2.9-3.6 mm long; corolla lobes 0.8-1.2
mm long, globose sessile glands present,
eglandular trichomes 2-8 per lobe. Achenes
narrowly ellipsoidal, 0.8-0.9 mm long, faintly
ribbed, dark brown, with 0-25 twin hairs each
<0.1 mm long, evenly distributed or mostly in
distal half; globose sessile glands at junction
between achene and pappus absent; pappus
2.1-3 mm long. Figs. 19, 20A-D.

Additional selected specimens examined : Australia:
Western Australia. S side of Cockburn Range, c. 13 km
W of King River, Jul 1974, Carr 3338 & Beauglehole
47116 (PERTH); Langi, 4 km S of Prior Point and
33 km SSW of Kuri Bay, Jul 1990, Kenneally 11055
(PERTH); Python Cliffs, Marigui Promontory, Prince
Regent River Reserve, Aug 1974, Kenneally 2138
(CANB, PERTH); The Grotto, Deception Ranges, near
Kimberley Research Station, Jul 1952, Lazarides 3071
(BRI, CANB, DNA); Mt Trafalgar, Prince Regent River
Reserve, Aug 1974, George 12698 (CANB, PERTH);
Inglis Gap, Gibb River Road, west Kimberley, Jun 1988,
Wilson 12959 (PERTH); Mirima [Hidden Valley] N.P,
Jun 1989, Kenneally 10947 (DNA, PERTH); Sunday
Island, Buccaneer Archipelago, Jun 1982, Kenneally
8250 (DNA, MEL, PERTH); Bungle Bungle N.P, Bull
Creek, Jun 1989, Cowie 874 (DNA, MEL, PERTH);
Gibb River Road, Home Valley Station, East Kimberley,
Jun 1992, Menkhorst 1067 (DNA, MEL, PERTH).
Northern Territory. South Bay, Bickerton Island, Gulf
of Carpentaria, Jun 1948, Specht 617 (A, BRI, CANB,
PERTH); Victoria River Gorge, May 2001, Leach &
Cowie 4659 (DNA); Gregory N.P, S of Victoria River
pub. May 2001, Risler & Smith 687 (DNA); Spirit Hills
Conservation Area; Nancy’s Gorge, Aug 1996, Cowie
7136 (DNA); 2 km S of Myra Falls, Jun 1993, Brennan
2355 (DNA); Fitzmaurice River, May 1994, Barritt 1270
(DNA, MEL); Keep River Gorge, Jun 1995, Egan 5004
(DNA); Bartelumba Bay, Groote Eylandt, Aug 1975,
Levitt 437 (CANB, DNA, NE, NT); Mt Brockman near
Koongarra Saddle, 2 km N of Koongarra, May 1980,

Lazarides 8905 (CANB, DNA); Nitmiluk N.P, eastern
boundary. May 2002, Michell 3877 (DNA); Gregory
N.P., c. 29 km NW of Bullita Outstation, Apr 1996, Coles
103 & Duretto (DNA, MEL); Keep River N.P, Apr 1982,
King 65 (DNA); Bauhinia Downs Station, Alligator
Stockyards, May 1985, Wightman 1835 & Leach
(CANB, DNA, NT); Limmen Bight, May 1996, Egan
5396 & Michell (DNA). Queensland. Burke District:
‘Westmoreland’, Big Amphitheatre, May 1997, Forster
BIF21039 & Booth (BRI, DNA).

Distribution and habitat : A widespread
species distributed in the northern Kimberley
of Western Australia, the ‘Top End’ of the
Northern Territory, and extending to the
north-west of Queensland (Map 9). The
occurrences of Pterocaulon tricholobum
are strongly associated with sandstone hills,
escarpments, scree slopes and gorges. The
vegetation type is variously recorded as
Triodia grassland, open savannah or shrubby
woodland. Soils are sandy.

Phenology : Flowers and fruits have been
recorded from May to September.

Affinities : Pterocaulon tricholobum is most
closely allied to P. globidiflorum , but differs
by the stem wings 2.8-5.5 mm wide (0.4-1.5
mm wide for P. globuliflorum ); the longest
inner bracts 4-4.8 mm long (3.2-3.5 mm
long for P. globuliflorum ); trichomes always
present on corolla lobes of hermaphrodite
floret (usually absent for P. globuliflorum );
achene hairs 0-25 (achene hairs 25-50 for
P. globuliflorum)., and pappus 2.1-3 mm long
(1.3-1.7 mm long for P. globuliflorum).

Notes : This species is distinctive in the
broad stem wings, rather small white globose
capitulescences that are often laterally and
the obvious trichomes on corolla lobes of
hermaphrodite floret.

The label of Coles 103 & Duretto records
that the plant “smells like Granny Smith
apples”.

Carr 3338 & Beauglehole 47116 is a mixed
gathering. All pieces on the CANB sheet are
Pterocaulon paradoxum ; both pieces on the
PERTH sheet are P. tricholobum.

Etymology : From the Greek trichos meaning
hair or thread, and lobus meaning lobes. This
is in reference to the conspicuous unicellular
hairs consistently borne on the corolla lobes of

322

Austrobaileya 8(3): 280-334 (2011)

Fig. 20. Pterocaulon tricholobum. A. hermaphrodite floret x 32. B. inner involucral bract (cilia not shown) x 16.
C. upper portion of inner involucral bract x 32, D. mature achene and basal portion of pappus x 32. Pterocaulon
verbascifolium. E. hermaphrodite floret x 32. F. inner involucral bract (cilia not shown) x 16. G. upper portion of inner
involucral bract x 32. H. mature achene and basal portion of pappus x 32. Pterocaulon xenicum. I. hermaphrodite
floret x 32. J. inner involucral bract (cilia not shown) x 16. K. upper portion of inner involucral bract x 32. L. female
floret x 32. M. mature achene and basal portion of pappus x 32. A-C from Egan 5004 (DNA); D from Cowie 7136
(DNA); E-H from von Oertzen 116 (DNA); I-L from Dunlop 1830 (DNA); M from George 14628 (PERTH).

Bean, Pterocaulon section Monenteles

323

the hermaphrodite floret. Other species have
this feature, but in this species the trichomes
are consistently present and obvious.

13. Pterocaulon verbascifolium (Benth.)
F.Muell., Syst. Census Austral PI. 79 (1882);
Monenteles verbascifolius Benth., FI. Austral.
3: 523 (1867). Type: Australia: Northern
Territory. Fitzmaurice River, s.dat., F.Mueller
s.n. (lecto: ¥312>2%6),fide Cabrera&Ragonese
(1978: 245).

Illustration : Cabrera & Ragonese (1978:
243).

Non-aromatic shrub 50-130 cm high; stems
very densely lanate; stem wings entire,
2.5-6(-8) mm wide. Leaves elliptical, ovate
or obovate, 37-70 x 20-43 mm, 1.6-2.3
times longer than wide, apex acute; margins
recurved, denticulate, with 5-24 pairs of
blunt teeth, sometimes obscure. Indumentum
comprising lanate multi-cellular hairs,
glandular hairs absent. Upper surface with
hairs erect to spreading, very dense, basal
cells short; globose sessile glands absent.
Lower surface with lanate hairs adpressed to
spreading, very dense, extending 0.5-0.9 mm
from the surface, basal cells not very short;
globose sessile glands sparsely present, but
hidden by hairs. Capitulescence spiciform
or cylindrical, 35-105 mm long and 17-30
mm wide at anthesis, 1.8-4.3 times longer
than wide; side branchlets never continuous,
at 30-90°; peduncles 3-40 mm long. Outer
involucral bracts 3.3-5.5 mm long, oblong
to oblong-spathulate, with dense spreading
lanate hairs on outer surfaces, inner surface
glabrous or lanate; glands absent; apex acute.
Inner bracts white on dorsal surface; longest
ones linear, 5.3-9.2 x 0.3-0.6 mm, apex acute,
upper margins with numerous small teeth;
cilia 12-30, up to 5.5 mm long, attached along
the proximal half and consistently extending
to the distal one third, not exceeding the
bract apex. Filiform (female) florets 25-58,
straw-coloured, corolla 37-7.5 mm long,
styles extending 0.4-0.9 mm beyond corolla.
Hermaphrodite floret 1(—2), corolla pink, 4-
6.5 mm long; corolla lobes 0.6-1 mm long,
globose sessile glands absent, eglandular
trichomes absent. Achenes narrowly
ellipsoidal, 1.1-1.3 mm long, terete or slightly

angular, dark brown, with 25-50 or more than
50 twin hairs each c. 0.1 mm long; globose
sessile glands at junction between achene
and pappus absent; pappus 3.5-6.9 mm long.

Figs. 20E-H, 21.

Additional specimens examined: Indonesia. Sumbawa,
s.dat., Zollinger 3421 (GH, L). Australia: Western
Australia. Dillie Gorge, Charnley River Station, Jul
2005, Byrne 1584 (PERTH); Carlton Hill Station, close to
Ningbing Site 3, Jun 2001, Handasyde & Start TH01137
(DNA, PERTH); Sir Graham Moore Island, Jul 1973,
Wilson 11317 (PERTH); SW from Mt Trafalgar, margin
of Saint George Basin, Jul 1990, Willing 212 (PERTH);
Fenelon Island, off Admiralty Gulf, Jul 1977, Kennea/ly
6412 (PERTH); Inglis Gap, King Leopold Range, May
1905, Fitzgerald 752 (PERTH); Bushfire Hill, Prince
RegentRiver Reserve, Aug 1974, George 12261 (PERTH);
around ‘Beverley Hills’ Homestead buildings, Jun 1992,
Mitchell 2557 (PERTH); Euro Gorge, Drysdale River
N.P, Aug 1975, Kenneally 4349 (PERTH). Northern
Territory. Aerodrome paddock, Manbulloo Station,
Apr 1971, Kruizinga s.n. (BRI, CANB); 7 miles NE of
Wilton River - Bulman Crossing, Jun 1972, Maconochie
1437 (BRI); ‘Bupa’, 4 km S Jarrong Yards, May 1994,
Leach 4269 & Walsh (BRI, DNA, MEL); Kakadu N.P,
Coronation Hill, Jul 1988, von Oertzen 116 (DNA);
Cave Creek Station, Snake Track, Jul 2002, Risler 1908
(DNA). Queensland, [unlocalised], s.dat.. Bowman
s.n. (K373287). Burke District: Gorge Creek gorge,
Bowthorn Station, 100 km W of Doomadgee, Aug 2001,
Addicott EPA1034 (BRI, CNS). Cook District: 21.1 km

5 of Georgetown, Aug 2010, Bean 29874 & McDonald
(BRI, CANB); 100 m from Georgetown - Forsayth
Road, 13 km S of Georgetown, Aug 2010, Bean 29861

6 McDonald (AD, BRI, NSW); 5.7 km E of Chillagoe,
Ramparts section, Chillagoe-Mungana Caves N.P, Jun
2011, McDonald 11687 & McDonald (BRI).

Distribution and habitat : Pterocaulon
verbascifolium is found commonly in the
northern Kimberley, including offshore
islands, and occasionally well inland. It is
widespread though sporadic in sub-coastal (but
not coastal) areas of the Northern Territory,
extending into northern Queensland, as far
east as Petford. Outside Australia, it is known
only from the island of Sumbawa in Indonesia
(Map 10). It inhabits lateritic plateaux or
rocky hill-slopes in eucalypt or Terminalia
dominated open woodland. Soil type varies,
and the geological substrate may be basalt or
sandstone.

Phenology : Flowers and fruits are recorded
between May and August.

Affinities : The species is not obviously allied
to any other.

324

Austrobaileya 8(3): 280-334 (2011)

STATE/DIST : NT/DG 14^5156"S 133°1035"E

CC07 Open shrub to 1.2m. Foliage grey green, densely hairy. Uncommon
in midhigh open woodland of Corymbia laiifolia and Erythrophleum
ddorostachys with Pierocaulon serrulalum and P. sphacealamm.

Risler. J.A. 1908 llJul2002

Fig. 21. Representative specimen of Pterocaulon verbascifolium ( Risler 1908 [DNA])

Bean, Pterocaulon section Monenteles

325

Notes : This species is distinctive in the dense
cylindrical capitulescences; corollas, achenes
and pappus longer than in other species; no
globose sessile glands on corolla lobes of
hermaphrodite floret; indumentum of leaves
extremely dense, obscuring surfaces; stem
wings broad and the foliage not aromatic.

Pterocaulon verbascifolium is remarkable for
the large variation between collections in the
sizes of the floral parts. In some collections, the
female florets (for instance) are only 3.7 mm
long, and in other collections they are up to
7.5 mm long. There are correlated differences
in other floral parts; however, there does not
appear to be any geographical or ecological
pattern to this variation.

There is a specimen of Pterocaulon
verbascifolium at K collected by E.M.
Bowman, with the locality given merely
as “Queensland”. It is of interest because
Bowman did not travel any further north or
west than the upper reaches of the Flinders
River, near present-day Hughenden (Blake
1955). Hughenden is more than 200 km beyond
the currently known extent of the species.

14. Pterocaulon xenicum A.R.Bean species
novaaffinisP.ra'veoaliscaulinisangustioribus,
numero majore flosculorum femineorum in
quoque capitulo, lobis brevioribus corollae
flosculorum hermaphroditorum et pappis
longioribus differens. Typus: Australia:
Western Australia. McLarty Hills, Great
Sandy Desert, 5 August 1977, A.S. George
14628 (holo: PERTH; iso: CANB).

Shrub 25-40 cm high; stems numerous,
densely lanate and sometimes with a few
globose sessile glands; stem wings entire,
0.3-0.5 mm wide. Leaves oblanceolate or
obovate, 18-52 x 7—17 mm, 2.6-47 times
longer than wide, apex obtuse or acute;
margins recurved, denticulate, with 10-21
pairs of blunt teeth. Indumentum comprising
lanate multi-cellular hairs, glandular hairs
absent. Upper surface moderately bullate;
hairs erect to spreading, sparse to dense,
basal cells short; globose sessile glands
absent. Lower surface with lanate hairs erect
to spreading, very dense, extending 0.4-0.7
mm from the surface, basal cells not very
short; globose sessile glands yellow and

shining, moderately dense, 3-8 diameters
apart, but usually obscured by indumentum.
Capitulescence globose to ellipsoidal, 13-18
mm long, 13-16 mm wide at anthesis, 1-1.3
times longer than wide; side branchlets never
continuous, at 30-80°; peduncles 0-14 mm
long. Outer involucral bracts 2.5-2.9 mm
long, spathulate, with dense spreading lanate
hairs throughout on the outer surface, glabrous
on the inner surface or with sparse hairs at
distal end; glands absent; apex lacerate to
fimbriate (fimbriae 0.1-0.35 mm long). Inner
bracts pink or white on dorsal surface; longest
ones linear, 4.8-5.6 x 0.25-0.4 mm; apex
acute, upper margins conspicuously toothed
or with small teeth; cilia 6-9, up to 3.8 mm
long, mostly attached in proximal half and
few extending to the distal one third. Filiform
(female) florets 26-44, pink, corolla 3.2-4.1
mm long, styles extending 0.2-0.4 mm beyond
corolla. Hermaphrodite floret solitary, corolla
pink, 3.5-4.3 mm long; corolla lobes 0.3-0.5
mm long, globose sessile glands present,
eglandular hairs absent or present. Achenes
narrowly ellipsoidal, 0.8-0.9 mm long, terete
or slightly angular, dark brown, with 25-50 or
>50 twin hairs each c. 0.05 mm long, evenly
distributed; globose sessile glands at junction
between achene and pappus sparsely present;
pappus 3.3-4.2 mm long. Figs. 20I-M, 22.

Additional specimens examined : Australia: Northern
Territory. 46 miles [74 km] WSW of The Granites, Aug
1970, Dunlop 1830 (DNA). Western Australia, just N
of Dragon Tree Soak, Great Sandy Desert, Aug 1977,
George 14748 (PERTH); 21.1 km ESE of Warrawagine
Station Homestead, Pilbara IBRA (Site PHYE07), Aug
2006, van Leeuwen et al. 0028 (BRI, PERTH); 13.3 km
W of Warrawagine Station Homestead, Pilbara IBRA
(Site PHYE06), Aug 2006, van Leeuwen et al. 0027
(BRI, PERTH); E of Cherrabun Station, Oct 1952, s.coll.
(PERTH536415).

Distribution and habitat: Known from a
few localities in northern Western Australia,
extending from Warrawagine Station to the
Tanami Desert in Northern Territory (Map 4).
At the type locality it occurs on open plains in
association with shrubs including Acacia spp.,
Senna spp., Solanum lasiophyllum Dunal
ex Poir. and Grevillea juncifolia Hook. The
understorey is dominated by spinifex ( Triodia
spp.)

326

Austrobaileya 8(3): 280-334 (2011)

hO/jrfYffsf Queensland Herbarium (BRI)

Pf'To com/ on xen /curt A . Boo/)
Del v&*2/U Zt>{/

PERTH 00541605

\Q-Oob

duplicates

rjicje^MT'D

FLORA OF THE KIMBERLEY REGION - W.A. HERBARIUM

’Pi-esocAvdo^ C£>C-) R NWAJt.

Signed _.. 1=^ 19^

WESTERN AUSTRALIAN HERBARIUM, PERTH
Flora of Western Australia

j£Lr~6 ^5 ~cv*; Arv-cSy .

‘f&A. / .

A t<J -4m*,

x*t /~<M .. Gx~j- S*~A*.

L 00 - A/.X}.

Co ll. <A. S.'^<rr^ - /x-6x§’ J- ^^, 19 ??

Fig. 22. Holotype of Pterocaulon xenicum (i George 14628 [PERTH]).

Bean, Pterocaulon section Monenteles

327

Phenology : Flowers and fruits are recorded
for August and October.

Affinities : The species shows affinity with
Pterocaulon sphacelatum and P. niveum.
Pterocaulon xenicum differs from P. niveum
by the stem wings 0.3-0.5 mm wide (0.5-1.5
mm for P. niveum ); the 26-44 female florets
(11-17 for P. niveum ); the corolla lobes of the
hermaphrodite floret 0.3-0.5 mm long (0.7-

1.1 forP. niveum ); and the pappus 3.3-4.2 mm
long (2-3.1 mm for P. niveum ).

Pterocaulon xenicum differs from
P. sphacelatum by the stem wings 0.3-0.5 mm
wide (0.7-2.0 mm forP. sphacelatum ); the longest
inner bracts 4.8-5.6 mm long (3.5-4.6 mm
for P. sphacelatum ); the female florets 3.2-4.1
mm long (2.4-3.2 mm for P. sphacelatum ); the
hermaphrodite florets 3.5-4.3 mm long (2.5-37
mm long for P. sphacelatum ); the pappus 3.3-

4.2 mm long (2.3-3.3 mm for P. sphacelatum );
and the outer bracts with lacerate or fimbriate
apices (acute apices for P. sphacelatum ).

Notes : This species is distinctive for the
stem wings being very narrow and almost
redundant; leaf indumentum very dense on
lower surface; corolla lobes of hermaphrodite
floret short; none or few cilia visible on
protruding part of inner bracts; apices of outer
bracts lacerate to fimbriate and the relatively
long hermaphrodite florets.

Etymology : Derived from the Greek word
xenos meaning ‘stranger, foreigner’ and the
suffix -icus ‘belonging to’. This alludes to the
specimens of this species found in amongst
collections of Pterocaulon sphacelatum , and
looking like ‘strangers’.

Postulated hybrids

Pterocaulon intermedium x p. paradoxum.

Specimens examined: Australia: Western Australia.
9 km N of Bunda Bunda Mill on Baldwin Creek
track, Dampier Peninsula, Jun 1984, Kenneally 9078
(PERTH).

This specimen has broad stem wings,
somewhat ellipsoidal capitulescences, achenes
c. 0.8 mm long and scarcely acute leaves. These
features do not fit Pterocaulon intermedium ,
nor do they quite fit P. paradoxum , although
the specimen has similarities to both. This

specimen is probably a hybrid between these
two species, both of which occur on the
Dampier Peninsula.

Pterocaulon serrulatum var. velutinum x
P. sphacelatum.

Specimens examined: Australia: Western Australia.
4.8 km E of Donkey Creek, on the Gibb River Road,
Sep 2006, Sweedman 6984 (PERTH); 6.5 km NE of
Mary River Crossing, Great Northern Highway, 92 km
SW of Halls Creek, Jun 1976, Beauglehole ACB53238
(PERTH); Site PHYE09, 10.3 km SW of Warrawagine
Station Homestead, Pilbara, Aug 2006, van Leeuwen et
al. 0035 (PERTH).

These specimens have glandular outer
involucral bracts as in Pterocaulon
serrulatum , but the leaves are smaller and
rather more bullate than is usual for that
species. Furthermore, the stem wings are
almost entire, with just the occasional tooth.

Pterocaulon ciliosum x p. serrulatum var.
serrulatum.

Specimens examined: Australia: Queensland. North
Kennedy District: 22.5 km (by road) SW of Herberton,
on the Silver Valley road. May 1983, Conn & Clarkson
1177 (BRI, CANB, DNA, NSW).

This specimen has narrower stem wings than
Pterocaulon serrulatum , and the toothing
is less than usual for that species. The
capitulescences are almost globose and the
protruding part of inner bracts has many long
cilia, both features of Pterocaulon ciliosum.

Excluded names

Pterocaulon tomentosus Boerl., Handl. FI.
Ned. Ind. (Boerlage) 2(1): 240 (1891).

This is a nomen nudum.

Monenteles tomentosus Sch. Bip., Syst. Verz.
(Zollinger) 122(1854).

This is a nomen nudum.

Gnaphalium redolens G.Forst., FI. Ins. Austr.
91 (1786).

This is a nomen nudum.

Acknowledgements

The Directors of several Herbaria have
kindly sent specimens on loan from their
institutions. I gratefully acknowledge
G.Rouhan (P), M.Pignal (P), P.Morat (P)

328

and P. Schafer (MPU) for their assistance or
advice regarding type specimens collected
from New Caledonia. I am grateful to N. Hind
(K) who dealt with my loan request promptly
and efficiently. V.Fonjallaz (G) and C.Nepi
(FI) kindly sent images of type specimens
held at those institutions. I thank G.von
Poser of Porto Alegre, Brazil, for sending a
reprint of a Pterocaulon chemistry paper.
K.McDonald, R.Booth and D.Kelman have
collected additional Pterocaulon material
at my request. W. Smith (BRI) provided the
illustrations of florets, bracts and achenes,
and PBostock (BRI) expertly translated the
diagnoses into Latin.

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330

Map 2. Distribution of Pterocaulon ciliosum ■. P. niveum O

Bean, Pterocaulon section Monenteles 331

Map 3. Distribution of Pterocaulon discolor ▲, P. globuliflorum □, P. sphaeranthoides •

Map 4. Distribution of Pterocaulon intermedium •, P. xenicum ▲

332

Austrobaileya 8(3): 280-334 (2011)

Map 5. Distribution of Pterocaulon paradoxum

Map 6. Distribution of Pterocaulon redolens

Bean, Pterocaulon section Monenteles

333

Map 7. Distribution of Pterocaulon serrulatum var. serrulatum •, P. serrulatum var. velutinum ▲

Map 8. Distribution of Pterocaulon sphacelatum

334

Austrobaileya 8(3): 280-334 (2011)

Map 9. Distribution of Pterocaulon tricholobum

Map 10. Distribution of Pterocaulon verbascifolium

Heliotropium microspermum E.J.Thomps.
(Boraginaceae), a new species from Queensland

E. J. Thompson

Summary

Thompson, E.J. (2011). Heliotropium microspermum E.J.Thomps. (Boraginaceae), a new species
from Queensland. Austrobaileya 8(3): 335-339. The new species Heliotropium microspermum
E.J.Thomps., endemic to Cape York Peninsula, Queensland, is described and illustrated.

Key Words: Boraginaceae, Heliotropium , Heliotropium microspermum , Queensland flora, taxonomy,
new species

E.J. Thompson, Queensland Herbarium, Department of Environment & Resource Management,
Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: john.
thompson@der m. qld. gov. au

Introduction

The genus Heliotropium L. comprises 84
species in Australia (Craven 1996, 2005a,b)
with 43 (38 native, five naturalised) occurring
in Queensland (Thompson 2010). A critical
review of Heliotropium specimens at the
Queensland Herbarium has revealed a new
species for Cape York Peninsula.

Materials and methods

Morphological data were collected from dried
herbarium material at BRI. Description of the
new species largely follows Craven’s (1996)
format.

Indumentum is a useful character in
Australian Heliotropium and is usually
“composed of unicellular, simple, non-
glandular hairs (the ‘standard’ hairs) with
multicellular, simple, glandular hairs also
occurring in some species. The walls of
standard hairs may be smooth or ornamented”
(Craven 1996). Craven (1996) refers to
indumentum as having more than one class of
hair when hairs can be differentiated, such as
long coarse hairs and shorter fine hairs.

Mericarps are essential for identifying
Heliotropium species and the mericarp
characters used were length, commissure (side
faces) pit shape and size; presence or absence
of a food body in the pits; size, distribution
and type of the hairs on the back; and shape
of the apex. Although Craven (1996) provided

Accepted for publication 23 August 2011

images of some mericarp indumentum types
he did not present illustrations of mericarps
for each species. Illustrations of mericarps
are provided here for the new species and its
putative allies. Other significant characters
included leaf width and hair type, inflorescence
characters, and plant growth form.

Taxonomy

Heliotropium microspermum E.J.Thomps.
species nova affinis H rhadinostachyum
Craven sed pilis applanatis fortuito dispositis,
fovea circulari minore in mericarpiis
minoribus et corolla breviore differens. Typus:
Queensland. Cook District: 12.3 km along
Pormpuraaw Road from Gulf Development
Road near Musgrave, 15 November 2010,
K.R.McDonald KRM10095 (holo: BRI).

Prostrate to ascending perennial herbs,
5-15 cm tall and to 15 cm broad. Branches
moderately hairy with ascending standard
hairs of one class. Adult leaves ascending,
linear to narrowly elliptic, 5-13 mm long, 2-3
mm wide, base attenuate, apex acute; petiole
short; abaxial surface moderately densely
hairy with appressed to ascending hairs;
adaxial surface with ornamented hairs, 07-1.2
mm long, similar to the abaxial surface only
hairs tending towards more numerous along
the midrib. Cyme on mature plants at anthesis
straight, simple or branched, bracteate; bracts
conspicuous, linear to narrowly elliptic, 2.5—
4.5 mm long, touching to separated up to one
length along the cymes, with ornamented
hairs on adaxial surface appressed on lamina

.i—----'-'T.

336

Austrobaileya 8(3): 335-339 (2011)

Fig. 1. Heliotropium microspermum. A. corolla *16. B. corolla expanded demonstrating anthers and appressed hairs
in throat x 16. C. pistil x 32. D. plant x 0.6. E. calyx x 16. F. mericarp back view x 40. G. mericarp side face x 40. All
from McDonald KRM10095 (BRI). Del. W.Smith.

Thompson, Heliotropium microspermum

337

and ascending on margin, abaxial surface
glabrous; varying from clearly different to
similar in shape and size to the leaves. Calyx
lobes with ornamented hairs 0.5-0.8 mm long
on the abaxial surface, appressed on the lamina
and ascending on the margin, adaxial surface
glabrous; outermost calyx lobe narrowly
ovate, 2-2.6 mm long, acute. Corolla 2.5-3
mm long, white, tube cylindrical, not swollen
around anthers at anthesis, 1.5-3 mm long, the
outer surface sparsely hairy with appressed
hairs, the inner surface glabrous in the throat;
lobes broadly triangular, 0.7-1 mm long, apex
acute. Anthers ovate, 0.3-0.5 mm long, apex
acuminate, papillate. Gynoecium c. 1.6 mm
long; ovary c. 0.5 mm long; style 0.5-0.7 mm
long; stigma 0.3-0.4 mm long. Mericarps
ovate to broadly ovate, 0.6-0.7 mm long, 0.4-
0.5 mm wide, 1.4 times as long as wide; apex
acute; outer surface with mostly appressed
to slightly ascending microhairs, 0.04-0.1
mm long, flattened, mostly distributed along
minute ridges over most of surface and
somewhat randomly orientated; commissures
with pit to about 14 width of commissure, the
pit elliptic, without a food body. Fig. 1.

Additional specimens examined: Queensland.
Cook District: 36.3 km from Dixie Station on track
to Killarney, Jun 1989, Clarkson 8154B & Neldner
(BRI); 13.9 km along Pormpuraaw Road from Gulf
Development Road, Nov 2010, McDonald KRM10093
(BRI); 6 km N of‘Fairview’ on the Peninsula Road, Apr
1980, Clarkson 3090 (BRI); 12.2 km along Pormpuraaw
Road from Cape York Development Road junction, Dec
2010, McDonald KRM10303 (BRI).

Distribution and habitat: Heliotropium
microspermum is only known from near
Musgrave on Cape York Peninsula (Map 1).
It has been found in woodland of Melaleuca
viridiflora Sol. ex Gaertn. on flat terrain,
and eucalypt woodland dominated by
Eucalyptus tetrodonta F.Muell. or Corymbia
stockeri subsp. peninsularis (K.D.Hill &
L.A.S.Johnson) A.R.Bean or C. nesophila
(Blakely) K.D.Hill & L.A.S.Johnson on
granite hills with sandy soil.

Phenology: Flowers and fruits have been
recorded in June and November.

Affinities: Heliotropium microspermum is
similar to H rhadinostachyum with respect
to growth habit, appearance of the cymose
inflorescences, and leaf and bract shape,
size, and distribution along the branches.

Heliotropium microspermum differs most
notably from H. rhadinostachyum and H.
cunninghamii Benth. by the size and shape
of the corollas and mericarps, as well as
indumentum type and cover (Table 1).

Notes: Juvenile leaves of Heliotropium
microspermum can be larger than adult leaves
and therefore more clearly different from
the bracts. Juvenile plants of this species
may have very short cymes. The corolla of
Heliotropium microspermum can be a little
swollen around the anthers at fruiting.

Conservation status: Heliotropium

microspermum is only known from near
Musgrave. Inappropriate burning regimes
present the most significant potential threat
to this species. Further field observation is
required to determine the species population
size and area of extent.

Etymology: The specific epithet is derived
from the Greek micro (small) and spermum
(seed) alluding to the relatively small
mericarps for this species.

Map 1. Distribution of Heliotropium microspermum in
Queensland

338 Austrobaileya 8(3): 335-339 (2011)

Table 1. Comparison of morphological characters for Heliotropium microspermum,

H. rhadinostachyum and H. cunninghamii

Character State

H. microspermum

H. rhadinostachyum

H. cunninghamii

mericarp length (mm)

0.6-0.7

0.9-1.2

0.6-1.2

mericarp apex shape

acute

rounded

acute

mericarp microhair
length (mm)

0.04-0.1

to c. 0.05 or
occasionally absent

0.05-0.1

mericarp microhair
type

flattened

papillate

circular in cross-
section

mericarp microhair
distribution

most of surface

upper 14

upper 14-14

mericarp pit size

small, up to Vi width
of commissure

large, c. 14 the width of
the commissure

small, up 14 width of
commissure

mericarp pit shape

elliptic

elliptic

circular

mericarp outer surface

minutely ridged over
most of surface

mostly smooth,
occasionally minutely
ridged in upper 14

smooth

corolla length (mm)

2.5-3

3-4

5-6

corolla tube shape at
anthesis

cylindrical

swollen around anthers

swollen around anthers

calyx lobe trichome
length (mm)

0.5-0.8

0.5-0.8

0.3-0.4

bract length (mm)

2.5-4.5

2-4.5

1-2.5

bract

conspicuous; leaves

conspicuous; leaves

inconspicuous; leaves

distinctly different to

distinctly different to

distinctly different

similar

similar

bract imbrication

touching to separated

overlapping by 14 a

widely separated

by about 1 length

length to separated by

up to 2 lengths

Acknowledgements

I am grateful to David Halford (BRI) and
Tony Bean (BRI) for reading drafts of the
manuscript. I thank Will Smith (BRI) for
the illustrations and map, and Peter Bostock
(BRI) for the Latin diagnosis.

References

Craven, L.R. (1996). A taxonomic revision of
Heliotropium (Boraginaceae) in Australia.
Australian Systematic Botany 9: 521-657.

- (2005a). Malesian and Australian Tournefortia

transferred to Heliotropium and notes on
delimitation of Boraginaceae. Blumea 50: 375-
381.

- (2005b). Seven new species of Heliotropium

(Boraginaceae) from the monsoon and arid
zones of Australia. The Beagle 21: 11-25.

Thompson, E. J. (2010). Boraginaceae. In P.D. Bostock &
A.E. Holland (eds.). Census of the Queensland
Flora 2010 , pp. 29-30. Queensland Herbarium,
Department of Environment & Resource
Management: Brisbane.

Thompson, Heliotropium microspermum

339

Fig. 2. A-D: Heliotropium rhadinostachyum. A. corolla * 16. B. outer calyx lobe x 16. C. mericarp backview x 40.
D. mericarp side view x 40. E-H: Heliotropium cunninghamii. E. corolla x 16. F. outer calyx lobe x 16. G.
mericarp back view x 40. H. mericarp side face x 40. A-D from Clarkson & Neldner 8146 (BRI); E-H from
Thompson & Booth TAN239 (BRI). Del. W.Smith.

Two new species of Pluchea Cass. (Asteraceae:
Plucheinae) from Queensland, Australia

A.R.Bean

Summary

Bean, A.R. (2011). Two new species of Pluchea Cass. (Asteraceae: Plucheinae ) from Queensland,
Australia. Austrobaileya 8(3): 340-346. The new species Pluchea punctata A.R.Bean and P.
xanthina A.R.Bean are described from Queensland, with the latter replacing the misapplied name
P. dioscoridis (L.) DC. Illustrations, a distribution map, and notes on the habitat and conservation
status are provided for each new species. A note is given regarding the apparent sexual dimorphism
of Pluchea baccharoides. A key to the identification of Queensland species of Pluchea is included.

Key Words: Asteraceae, Pluchea, Pluchea baccharoides, Pluchea punctata, Pluchea xanthina,
Australia flora, Queensland flora, taxonomy, new species, identification key

A.R.Bean, Queensland Herbarium, Department of Environment & Resource Management, Brisbane
Botanic Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia. E-mail: tony.bean@
derm.qld.gov.au

Introduction

Pluchea Cass, is a genus of about 50 species,
with representatives throughout the tropics
and subtropics of the world, but especially
in eastern Africa, the Arabian Peninsula and
north-western Australia (King-Jones 2001;
Thulin 2004). The Australian members of
the genus are perennial shrubs or subshrubs,
often woody, with alternate toothed leaves,
and pink tubular florets. The involucral bracts
are acute, and borne in several rows. The
achenes are cylindrical to ellipsoidal, and the
pappus comprises many barbellate capillary
bristles.

Hunger (1996) described the new species
Pluchea dunlopii from Australian material,
and soon after provided a revision of the
genus for Australia wherein six species were
enumerated (Hunger 1997).

Two new species are described here, both
endemic to Queensland. Pluchea xanthina
A.R.Bean is a moderately widespread species
for which the name P. dioscoridis (L.) DC.
has been misapplied. Pluchea punctata
A.R.Bean is a newly discovered species with
a very restricted distribution. Both species
are illustrated and compared with related
taxa. A distribution map is provided, as is an
identification key for the Queensland species
of Pluchea.

Materials and methods

This paper is based on a study of herbarium
specimens at BRI, an image of the type
specimen of Blumea dioscoridis at LINN
and examination of Pluchea populations and
plants in the field. Measurements of florets
were made from material preserved in spirit
or reconstituted in boiling water, while leaves,
bracts and achenes were measured from dried
specimens.

Abbreviations used in the text include N.R
(National Park) and S.F. (State Forest).

Taxonomy

Pluchea xanthina A.R.Bean species nova

affinis P. wallichianae, sed foliis petiolatis
angustioribus, bracteis involucral ibus
exterioribus longioribus, marginibus
superioribus integris in bracteis involucral ibus
omnibus et corollae lobis longioribus
flosculorum discoideorum differens. Typus:
Queensland. South Kennedy District:
Llanarth Back Range Road, 9.4 km S of
junction with Scartwater Road, 15 May 1991,
VJ.Neldner 3479 & E.J.Thompson (holo:
BRI; iso: AD, K).

Pluchea dioscoridis auct. non (L.) DC.;
Mueller (1889); Bailey (1900); Hnatiuk (1990);
Henderson (1997); Holland (2007, 2010).

Accepted for publication 27 June 2011

Beam. Two new Pluchea (Asteraceae)

341

Woody shrub to 150 cm high, rarely higher.
Stems greento brown, terete, erectto spreading,
well-branched, with sessile yellow glands and
multicellular eglandular hairs to 0.3 mm long,
dense on young stems, glabrescent on older
stems. Leaves petiolate, narrowly-elliptic to
oblanceolate, spreading, not decurrent, 24-68
x 4-17 mm, 3.8-6.4 times longer than broad,
margins denticulate or occasionally dentate,
apex acute; both surfaces with many sessile
yellow glands and multicellular eglandular
hairs; petioles 5-15 mm long. Capitula in
small terminal clusters, campanulate, 7-8 x
6-7 mm; peduncles 1—6(—16) mm long, with
1-3 lanceolate bracts near the apex, each l-3(-
9) mm long. Involucral bracts in several rows;
outer bracts ovate, 3.2-4.2 x 1.3—1.7 mm, with
apex acute, outer surface with sessile yellow
glands and dense eglandular hairs, margins
entire distally; median bracts elliptic, 4.7-5 x
1.2-1.6 mm, apex acute, distal half of outer
surface with sessile yellow glands and dense
eglandular hairs, upper margins entire; inner
bracts linear, 6-6.7 x 0.4-0.8 mm, apex acute,
outer surface glabrous, upper margins entire.
Receptacle epaleate, verrucose, flat, 1.6-2.8
mm diameter. Outer florets 75-100, female;
corolla very slender, 4.3-6 mm long, pink;
lobes 3, triangular, 0.2-0.3 mm long, glabrous;
style bifid, exserted, glabrous; pappus 4.5-5.1
mm long, pappus bristles 9-13, all of similar
length, barbellate. Disc florets 4-8, each with
an ovary and divided style but functionally
male; corolla cylindrical, 4.5-6 mm long,
purple; lobes 5, triangular, 0.7-1 mm long,
glandular; anthers c. 2 mm long, including tail
c. 0.2 mm long; style bifid, exserted by 0.5-1
mm, with acute sweeping hairs; pappus 5.2-6
mm long, pappus bristles c. 20, barbellate, all
of similar length. Achenes cylindrical, 0.9-1.2
mm long, with scattered appressed twin hairs;
carpopodium prominent, white. Fig. 1.

Additional selected specimens examined : Queensland.

Burke District: Micky Spring, ‘Clyde Park’, N of
Hughenden, Aug 1984, Godwin C2715 (BRI). North
Kennedy District: Lynd Scrub, c. 64 km SW to SSW of
Mt Garnet, Aug 1948, Smith 3857 (BRI); 15 km S of ‘Mt
McConnell’ Homestead, Jun 1992, Thompson CHA60
& Sharpe (BRI). South Kennedy District: 6 km NE of
‘Moonoomoo’ Homestead in Darkies Range, Apr 1992,
Thompson BUC354 & Simon (BRI); Cudmore N.P.,
Sep 2001, Augustyn JA5 (BRI). Leichhardt District:
Gogango Range, near Edungalba, Sep 1943, Blake 15354

(BRI); Southern end of S.F. 236, SW of Blackwater, Nov
2002, Bean 19528 (BRI); c. 30 miles [48 km] WSW of
Springsure, Sep 1966, Everist 7939 (BRI); Goodliffe
section, Carnarvon N.P, 12 km W of eastern boundary
of park, along road from Springsure, Sep 2009, Halford
Q9690 (BRI); 17 miles [28 km] E of Duaringa, Aug
1963, Speck 1667 (BRI, CANB, NSW); ‘Early Storms’,
Carnarvon Creek, Sep 1974, Gittins 2764 (BRI,
NSW). Port Curtis District: 12.9 km by road SE of
Maryborough towards Rockhampton, Sep 1975, Coveny
6908 & Hind (BRI); ‘Maryvale’, between Byfield and
Yeppoon, Sep 1931, White 8036 (BRI); N of The Caves,
near Yaamba, Rockhampton, Aug 1992, Barton 4 (BRI).
Burnett District: Cole’s Lookout, Coominglah S.F., W
of Monto, Aug 1995, Bean 8856 (BRI, MEL).

Distribution and habitat: Pluchea xanthina is
endemic to Queensland. It occurs from Forty
Mile Scrub N.P. in northern Queensland south
to the Carnarvon Ranges and Monto, with
several collections from the Rockhampton
area (Map 1). The species grows on hilly
terrain in eucalypt woodland, often with
Corymbia citriodora (Hook.) K.D.Hill &
L.A.S. Johnson ox Eucalyptus crebra F.Muell.,
on sandy-loamto clay-loam soils. It sometimes
grows on the margins of brigalow forest, and
near Rockhampton it has also been recorded
growing on serpentinite.

Phenology: Flowers and fruits have been
recorded from May to November.

Notes: Mueller (1889) was the first to apply
the name Pluchea dioscoridis to Australian
material. This name was then employed by
Bailey (1900), and subsequent authors have
followed that usage. A high quality image
of the type of Blumea dioscoridis L. (LINN
992.6) has been viewed on the Linnean
Society website (Anon. 2010). It shows a rather
broad-leaved plant with sessile auriculate-
based leaves, clearly different to the taxon
found in Queensland. According to Boulos
(2002), the type was collected in Egypt by
F. Hasselquist. King-Jones (2001) recorded
Pluchea dioscoridis from central and northern
Africa and the Arabian Peninsula.

Pluchea xanthina differs from
P. dioscoridis in several characters based
on the description of the latter by King-
Jones (2001): leaf base attenuate (auriculate
in P. dioscoridis ); capitula 7-8 mm long
(4.5-(5-7)-9) mm in P. dioscoridis );
involucral bracts with margins entire distally

342

Austrobaileya 8(3): 340-346 (2011)

Fig. 1. Pluchea xanthina. A. flowering branchlet * 0.5. B. portion of leaf surface showing eglandular hairs and sessile
glands x 12. C. capitulum x 6. D. median involucral bract x 12. E. mature achene and pappus x 12. F. receptacle of
spent capitulum and bract-bases x 12. G. disc floret x 8. A, B from Neldner 3479 & Thompson (BRI); C, D from
Halford Q9690 (BRI); E-G from Bean 20718 (BRI).

Beam. Two new Pluchea (Asteraceae)

343

(fimbriate in P. dioscoridis)', inner involucral
bracts 6-6.7 mm long (4.2-5.2 mm long in
P. dioscoridis ); outer florets 4.3-6 mm long
(2.7-4 mm in P. dioscoridis ); and pappus
bristles all the same length (of varying length
in P. dioscoridis ).

Pluchea xanthina is perhaps most closely
allied to P. wallichiana DC., a species from
India and Pakistan. Pluchea xanthina differs
by the leaves 0.4-1.7 cm wide (1.5—2.5(—3) cm
in P. wallichiana ); leaves consistently petiolate
(consistently sessile and sometimes auriculate
in P. wallichiana ); outer involucral bracts 3.2-
4.2 mm long (2-2.5(-2.8) in P. wallichiana );
involucral bract margins entire (fimbriate in
P. wallichiana ); and corolla-lobes of the disc
florets 0.7-1 mm long (0.4-0.6 mm long in
P. wallichiana).

Conservation status : A widespread species,
that it is not currently under threat.

Etymology : The epithet is from the Greek
xanthos meaning yellow and inos, of the
nature of It refers to the shining yellow glands
that are found commonly on many parts of the
plant.

Pluchea punctata A.R.Bean species nova

affinis P. baccharoidi sed capitulo longiore,
marginibus glandularibus bractearum
involucralium, flosculis discoideis corollae
lobis quattuor, pappo longiore et acheniis
ellipsoidalibus brevioribus differens. Typus:
Queensland. Cook District: Gilbert River
Holding, 78 km by road S of Forsayth,
13 August 2010, A.R.Bean 29877 &
K.R.McDonald (holo: BRI [1 sheet + spirit];
iso: AD, B, CNS, MEL, NSW, US).

Woody shrub to 80 cm high. Stems green
to brown, terete, erect to spreading, well-
branched, glabrous, viscid. Leaves sessile,
1 inearto very narrowly oblanc eolate, spreading,
not decurrent, 18-54 x 1-2.5 mm, 13-23 times
longer than broad, viscid or resinous, margins
entire, apex acute; both surfaces punctate
with conspicuous embedded glands. Capitula
in small terminal clusters, narrowly ovoid to
campanulate, 7-10 x 4-5 mm; peduncles 1-10
mm long, with 1-3 lanceolate bracts near the
apex, each 1.5-2.5 mm long. Involucral bracts
in several rows; outer bracts ovate, 2.5-3.5 x

1.5-1.7 mm, apex acuminate, outer surface
and margins with stipitate glandular hairs;
median bracts elliptic, 3.5-47 x 1.2-1.8 mm,
apex obtuse to acuminate, outer surface and
distal margins with stipitate glandular hairs;
inner bracts linear, 5-6.3 x 0.4-0.6 mm, apex
acute, glabrous or with a few stipitate glands
near apex. Receptacle epaleate, verrucose,
flat, c. 2 mm diameter. Outer florets 80-120,
female; corolla very slender, 4.2-5.5 mm
long, pink; lobes 3, triangular, 0.15-0.25 mm
long, glabrous; style bifid, exserted, glabrous;
pappus 3.7-5 mm long, pappus bristles 13-17,
all of similar length, barbellate. Disc florets
8-12, each with an ovary and divided style
and appearing bisexual but functionally male;
corolla cylindrical, 5.2-6.5 mm long, purple;
lobes 4, triangular, each 0.6-0.7 mm long,
glandular; anthers c. 2 mm long, including
a tail c. 0.3 mm long; style bifid, exserted
by 1.5-2 mm, with acute sweeping hairs;
pappus 4.3-5.5 mm long, pappus bristles c.
15, all of similar length, barbellate. Achenes
narrowly ellipsoid, 0.8-1.0 mm long, with
scattered appressed twin hairs; carpopodium
prominent, white. Fig. 2.

Additional specimen examined: Queensland. Cook
District: Gilbert River holding, 40 km by road from
Cobbold Gorge turnoff, Oct 2007, McDonald KRM6961
(BRI, CANB).

Distribution and habitat: Pluchea punctata
is known only from the type locality (Map
1). It occurs on the lower slope of a steep hill
amongst Triodia sp., in an exposed area with
scattered trees of Eucalyptus microneura
Maiden & Blakely and Grevillea striata R.Br.
Soil is a reddish-brown sandy-loam.

Phenology: Flowers and fruits have been
recorded in August and October.

Affinities: Pluchea punctata is superficially
very similar to P. baccharoides (F.Muell.)
Benth. as both are small woody shrubs with
viscid gland-dotted, linear leaves. However,
Pluchea punctata can be distinguished by
the longer leaves 18-54 mm long (6.5-19
mm long for P. baccharoides ); capitula 7-10
mm long (6-7 mm long for P. baccharoides );
involucral bracts with stipitate glandular hairs
on margins (eglandular fimbriate margins
for P. baccharoides ); disc florets 8-12 (1-2
or 13-18 for P. baccharoides ); disc florets

344

Austrobaileya 8(3): 340-346 (2011)

Fig. 2. Pluchea punctata. A. flowering branchlet x 0.5. B. portion of leaf surface showing embedded glands x 16.
C. capitulum x 6. D. median involucral bract x 16. E. disc floret x 8. F. mature achene and pappus x 16. A-E from Bean
29877 & McDonald (BRI); F from McDonald KRM6961 (BRI).

Beam. Two new Pluchea (Asteraceae)

345

with 4 glandular corolla lobes (5 eglandular
lobes for P. baccharoides ); female florets
with pappus 3.7-5 mm long (2.5-3 mm for
P. baccharoides ); and achenes narrowly
ellipsoid, 0.8-1 mm long (linear, 1.4-1.7 mm
long for P. baccharoides ). The hillside habitat
of Pluchea punctata is quite different to the
strictly riparian habitat of P. baccharoides.

Conservation status : There are between 40
and 50 plants at the type locality. Similar
habitats along the roadside north and
south of the type locality were searched
without locating any additional plants. A
conservation status of Critically Endangered
is recommended, based on criterion D of the
IUCN Red List (IUCN 2001).

Etymology : The epithet is from the Latin
punctatus , dotted, which derives from the
Latin punctum ‘a spot, dot, or point’, and
refers to the dot-like glands embedded in the
leaves of this species. Such glands are found
in only a few species of Pluchea.

A note on Pluchea baccharoides

Hunger (1997) noted that Pluchea
baccharoides produces both homogamous
and heterogamous capitula; however, she gave
no indication as to whether homogamous and
heterogamous capitula appear on the one plant
or on different plants. Hunger (1997) recorded
2-30 disc florets and 0-60 female florets
for P. baccharoides ; these observations are
correct but rather misleading, as they imply a
continuum of floret numbers.

There are currently 17 flowering specimens
of Pluchea baccharoides at BRI. Of these,
11 were found to have only heterogamous
capitula with 40-90 female florets and
1 or 2 functionally-male disc florets (not
“hermaphroditic”, as stated by Hunger). The
other six specimens bear only homogamous
capitula in which all 13-18 disc florets are
functionally male, and female florets are
lacking. The absence, in this sample, of any
individual specimen with both homogamous
and heterogamous capitula suggests that the
species is subdioecious.

Key to Queensland species of Pluchea

1 Leaves and stems resinous or viscid.2

1. Leaves and stems neither resinous nor viscid.4

2 Leaves broadly obovate, toothed; hairs (0.1-0.25 mm long) frequent on

stems and leaves.P. dunlopii

2. Leaves linear, entire; hairs absent on stems and leaves.3

3 Margins of involucral bracts fimbriate, eglandular; disc florets with five

corolla lobes; achenes 1.4-1.7 mm long.P. baccharoides

3. Margins of involucral bracts with stalked glandular hairs; disc florets

with four corolla lobes; achenes 0.8-1 mm long.P. punctata

4 Leaves and stems with surfaces obscured by a very dense indumentum

P. ferdinandi-muelleri

4. Leaves and stems glabrous or hairy but surfaces readily visible

5 Leaves and stems with eglandular septate hairs.6

5. Leaves glabrous or with stalked glandular hairs, but eglandular septate hairs absent.... 7

6 Leaves and stems with many yellow sessile glands.P. xanthina

6. Leaves and stems without sessile glands.P. indica

7 Leaf-bases not decurrent; stems and leaves with many stalked glandular

hairs; inner involucral bracts 6-8 mm long.P. dentex

7. Leaf-bases decurrent on stems; stems and leaves glabrous; inner involucral

bracts 3.5-5 mm long.P. rubelliflora

346

Austrobaileya 8(3): 340-346 (2011)

Acknowledgements

I am grateful to Keith McDonald for taking
me to the population of Plucheapunctata that
he discovered. My thanks are also due to Will
Smith for the illustrations and Peter Bostock
for the Latin diagnoses.

References

Anonymous. (2010). Linnean Herbarium Online. The
Linnean Society of London: Piccadilly, United
Kingdom, http://linnean.org/ [Accessed 12 Nov.
2010 ],

Bailey, F.M. (1900). The Queensland Flora, Part III. J.
Diddams & Co.: Brisbane.

Boulos, L. (2002). Flora of Egypt , Volume 3
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(Compositae, Plucheeae) from Australia.
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(Compositae, Plucheeae) in the Old World.
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22: 659-666.

Eucalyptus melanophloia subsp. nana D.Nicolle & Kleinig, a new
mallee ironbark (E. series Siderophloiae Blakely; Myrtaceae)
from central Australia and north western Queensland

D.Nicolle 1 * & D.A.Kleinig 2
Summary

Nicolle, D. & Kleinig, D. (2011). Eucalyptus melanophloia subsp. nana D.Nicolle & Kleinig, a new
mallee ironbark (E. series Siderophloiae Blakely; Myrtaceae) from central Australia and north western
Queensland. Austrobaileya 8(3): 347-355. The new subspecies Eucalyptus melanophloia subsp.
nana is described. The subspecies is of widespread but very scattered distribution in arid regions on
rocky sites from the Yaripilangu Range in the Northern Territory in the west (about 250 km WNW of
Alice Springs), eastwards to the Mount Isa area in north western Queensland. It is distinguished from
subsp. melanophloia by its consistently multi-stemmed, bushy mallee habit and this is unique among
the ironbark eucalypts ( Eucalyptus series Siderophloiae and E. series Rhodoxylori). Illustrations of
the new subspecies (including the holotype), a key to the subspecies of Eucalyptus melanophloia and
a distribution map are provided.

Key Words: Myrtaceae, Eucalyptus, Eucalyptus melanophloia , Eucalyptus melanophloia subsp.
nana , Australia flora. Northern Territory flora, Queensland flora, taxonomy, new subspecies,
ironbark, mallee

‘D.Nicolle, Currency Creek Arboretum, RO. Box 808, Melrose Park, South Australia 5039, Australia.
Email: dn@dn.com.au *author for correspondence

2 D. A.Kleinig, P.O. Box 395, Clare, South Australia 5453, Australia. Email: dendros@bigpond.com

Introduction

Eucalyptus melanophloia F.Muell. belongs
to E. series Siderophloiae Blakely (which
includes all the ironbark taxa in which
all the stamens are fertile - about 22
species) subseries Jugatae Blakely (only
E. melanophloia and E. shirleyi Maiden).
The subseries is distinguished within the
genus by the combination of the following
characteristics (amended from Brooker
2000 ):

Eucalyptus subgenus Symphyomyrtus
(Schauer) Brooker-Cotyledons folded in
seeds; buds bi-operculate; seeds with ventral
or terminal hilum; seed coat formed from
both integuments.

Eucalyptus section Adnataria L.D.Pryor
& L.A.S.Johnson ex Brooker-Cotyledons
reniform; seeds with a ventral hilum; leaf
venation usually densely reticulate; anthers
not versatile.

Accepted for publication 29 June 2011

Eucalyptus subsection Apicales Brooker -
Anthers erect at tip of filament; inflorescences
commonly terminal.

Eucalyptus series Siderophloiae —
Ironbarks; outer operculum shed early;
all stamens fertile, irregularly flexed.

Eucalyptus subseries Jugatae- Juvenile
leaves sessile; mature crown of juvenile
leaves.

Eucalyptus subseries Jugatae (with the
two species E. melanophloia and E. shirleyi )
was recognised by Brooker (2000) on the
basis of the sessile juvenile leaves forming
the mature crown, which distinguish the
subseries from E. subseries Subglaucae
Blakely (the remaining 20 species of E. series
Siderophloiae ), which have petiolate juvenile
leaves and a crown of adult leaves. However,
the distinction between these subseries is
weak and perhaps not representative of the
phylogeny of the group; indeed Eucalyptus
whitei Maiden & Blakely of E. subser.
Subglaucae forms substantial intergrading
populations with E. melanophloia of
E. subseries Jugatae (Holman et al. 2011).

348

Nevertheless, within Eucalyptus subseries
Jugatae , E. melanophloia is distinguished
from E. shirleyi in its less straggly habit,
smaller juvenile leaves, the occasional
development of intermediate or adult leaves
in some individuals (E. shirleyi is not known
to develop intermediate or adult leaves) and
the much smaller flower buds and fruits.
Interestingly, Eucalyptus shirleyi is quite
variable and may be composed of more than
one taxon. The distribution of Eucalyptus
subseries Jugatae can be roughly defined by
the distribution of E. melanophloia , with the
distribution of E. shirleyi (inland of Mackay
north to the Atherton Tableland region)
occurring largely within the northern part of
the distribution of E. melanophloia.

The new taxon described here was first
collected south of Mt Isa in Queensland as early
as 1951 and has subsequently been collected
numerous times south of Mt Isa on the Mt
Isa to Dajarra road, as well as occasionally
from elsewhere on the hills around Mt Isa.
The population on the Mt Isa to Dajarra road
has become well known because it is atypical
compared to Eucalyptus melanophloia
elsewhere in Queensland. Brooker & Kleinig
(1994: 347) mentioned the population in the
text .. occurring as a small mallee south of
Mt Isa”. Brooker & Kleinig (1994) included
an image of both the tree variant (subsp.
melanophloia) and the mallee variant (subsp.
nana) of E. melanophloia. Field and herbarium
studies in conjunction with cultivation trials
in a common garden indicate that the Mt Isa
populations are sufficiently distinct to warrant
taxonomic recognition.

The new subspecies was not collected in
central Australia (Northern Territory) until the
1990s, when Peter Latz collected specimens
from two widely separated localities
(Ennugan Mountains and Yaripilangu
Range). A number of other central Australian
collections have since been made in localities
in between, including at the type locality at
Mt Leichhardt.

Taxonomic recognition of the two variants
of Eucalyptus melanophloia is considered
appropriate due to the conspicuous distinction
in habit between the two geographically

Austrobaileya 8(3): 347-355 (2011)

disjunct variants. This is consistent with
the taxonomic recognition of other tree
versus mallee variants (genetically and not
environmentally determined) within the
eucalypts, such as Eucalyptus loxophleba
Benth. subsp. supralaevis L.A.S.Johnson
& K.D.Hill (tree) and subsp. lissophloia
L.A.S.Johnson & K.D.Hill (mallee).

The new subspecies ( Eucalyptus
melanophloia subsp. nana) has great
horticultural potential in arid regions, because
of its depauperate habit (for a eucalypt), crown
of silvery leaves, and habitat of rocky sites
(i.e. confined soil pockets) in a hot and dry
region.

The relatively recent discovery of this
new subspecies in Central Australia must be
associated with the remoteness and relative
paucity of botanical collections in the area
in which it occurs. The subspecies is not
rare (it can locally dominate some vegetation
communities) and it is conspicuous in the
field.

Material and methods

This paper is based on examination of
herbarium collection at the herbaria AD,
BRI, CANB and NSW, together with visits
to populations in the Northern Territory and
Queensland.

Taxonomy

Eucalyptus melanophloia F.Muell., J. Proc.
Linn. Soc., Bot. 3: 93 (1859). Type citation:
A tractu montano Newcastle Range usque ad
sinum Morton Bay Eucalypto crebra abunde
adsociata, solum sterilibus indicans’. Type:
Queensland: Dawson River, s.dat. [1856],
F.Mueller s.n. (lecto: K, material on upper
half of sheet, fide Bean [2010: 472]).

Bark rough throughout (to branches less
than 25 mm in diameter), moderately to
deeply fissured and hard (ironbark), dark
grey to black. Branchlets terete, waxy on
current season’s growth (wax eroded and
not apparent on old branchlets), lacking pith
glands. Seedling leaves opposite, sessile or
on petiole to 2 mm long, elliptical to ovate,
concolorous to slightly discolorous, dull,
greyish, usually waxy, 35-60 mm long,

Nicolle & Kleinig, Eucalyptus melanophloia

16-35 mm wide. Juvenile leaves opposite,
sessile, (cordate at base) broadly elliptical
to ovate (rounded to slightly pointed at tip),
concolorous, dull, greyish, waxy on current
season’s growth, persisting on reproductively
mature plants. Inflorescences held erect,
terminal or sometimes sub-terminal (growing
tip continuing). Peduncles terete or nearly so.
Pedicels terete. Flower buds with operculum
scar (but outer operculum sometimes still
present at tip at anthesis), diamond-shaped
to broadly fusiform, broadest below or at
hypanthium-operculum join, 6-9 x 2.8-3.2
mm, waxy; operculum bluntly conical to

349

conical, 3-5 mm long. Flower stamens cream,
all anthers fertile. Fruits waxy when young
(wax eroded and not apparent on old fruits).
Seeds compressed ovoid, 1-2 mm long, dark
grey-brown to black, finely reticulate; chaff
smaller, linear, orange brown.

Two geographically disjunct subspecies
are recognised in Eucalyptus melanophloia ,
differing most conspicuously and consistently
in habit but also generally differing (with
some degree of overlap) in leaf spacing and
size, and in fruit size.

Key to subspecies of Eucalyptus melanophloia

Tree, usually single-stemmed, 7 to 18 m tall; most internodes 20 to 50
mm long (central and eastern Queensland and northern New South Wales)

.subsp. melanophloia

Mallee, multi-stemmed, 2 to 6 m tall; most internodes 10 to 30 mm long
(Northern Territory and north western Queensland).subsp. nana

Eucalyptus melanophloia subsp.
melanophloia

Tree, usually single-stemmed at ground level,
7-18 metres tall; forming an inconspicuous
lignotuber. Juvenile leaves well-spaced
(internodes usually 20-50 mm). Juvenile
leaves 35-70 mm long, 22-48 mm wide.
Intermediate and adult leaves sometimes
produced in crown, 75-110 mm long, 13-30
mm wide. Inflorescences 7 or 9-flowered.;
peduncles 5-14 mm long; pedicels 1-5 mm
long. Fruits cupular to truncate-globose to
pyriform, 5-6 mm long, 5-6.5 mm wide, disc
descending then level to slightly ascending,
valves 3, 4 or 5, enclosed to slightly exserted.

Distribution and habitat : Widespread
and relatively common in north-eastern
Australia, from Narromine in New South
Wales northwards to west of Musgrave on
central Cape York Peninsula in Queensland
(Clarkson 2009), and inland in Queensland
to near Adavale and Croydon. A common
component of open woodland vegetation,
where it occurs on level to undulating
topography as well as on hills and ridges, in
skeletal or sandy soils (occasionally including

deep red sands) to well-drained loams.
Numerous woodland eucalypt species have
been recorded as associates, and although
the subspecies sometimes forms pure stands,
it is more commonly co-dominant with one
of a number of Eucalyptus section Adnataria
(ironbark or box) species, especially E. crebra
F.Muell.

Notes: Putative sporadic hybrids are known
with a number of other species, including
intra-serial hybrids with Eucalyptus crebra
(e.g. Forster 16089, 16229, 13104, 16281; see
Brophy et al. 2009), E. cullenii Cambage,
E. decorticans (F.M.Bailey) Maiden and

E. drepanophylla F.Muell. ex Benth. (in the
broad sense when including E. xanthoclada
Brooker & A.R.Bean) and inter-serial
hybrids with E. coolabah Blakely & Jacobs
and E. leptophleba F. Muell. (both E. series
Aquilonares Brooker) and E. populnea

F. Muell. and E. orgadophila Maiden & Blakely
(both E. series Buxeales Blakely). Extensive
intergrading populations between Eucalyptus
melanophloia subsp. melanophloia and
E.whitei are known where their distributions
adjoin in the Pentland to Jericho area of
Queensland (Holman et al. 2011).

350

Eucalyptus melanophloia subsp. nana
D.Nicolle & Kleinig subspecies nova a
subspecie typica habitu minore et pluricauli
“mallee”, foliis juvenilibus minoribus et
internodis brevioribus differt. Typus:
Northern Territory. Western slope of Mt
Leichhardt, 1 July 2008, D.Nicolle 5205 &
M.E.French (holo: AD; iso: CANB, DNA,
PERTH).

Eucalyptus melanophloia subsp. (Dajarra V. J.
Neldner 1523); Bean et al. (2010: 119).

Mallee, several-stemmed from ground
level, 2-6 metres tall; forming a lignotuber.
Juvenile leaves crowded (internodes mostly
10-30 mm). Juvenile leaves 18-55 mm long,
12-40 mm wide. Reproductively mature in
juvenile leaf phase, intermediate and adult
leaves rarely produced. Intermediate leaves
rarely produced. Adult leaves not known.
Inflorescences 7-11-flowered. Peduncles 5-
10 mm long. Pedicels 1-2 mm long. Fruits
cupular, truncate-globose, barrel-shaped
or slightly pyriform, 5-6 mm long, 4.5—5
mm wide, disc descending, valves 3 or 4,
enclosed. Fig. 1-3.

Additional selected specimens examined : Northern
Territory. 4 km NE of Mt Leichhardt, 30 km NNE of
‘Coniston’ Homestead, Sep 2006, Dugnid 22376 & Latz
(CANB, NT); Mt Leichhardt, 29 km NNE of ‘Coniston’
Homestead, Sep 2006, Duguid 22380 & Latz (CANB,
NT); Mt Leichhardt summit, Jul 2008, Nicolle 5207 &
French (AD); Mt Leichhardt, upper S slope, Aug 2005,
Albrecht 11677 & Latz (BRI, NT); Mt Leichhardt, lower
S-slope, Aug 2005, Albrecht 11660 & Latz (BRI, DNA,
NSW, NT); Mt Denison, 23 km NNW of ‘Coniston’
Homestead, Sep 2005, Latz 21028 (BRI, NT); SE end of
Yindjirbi Range, Coniston Station, Jul 2008, Nicolle 5203
& French (AD, CANB, DNA); Ti Tree Station, Ennugan
Mountains, Jul 1992, Latz 12366 (AD, DNA, NT); 13 km
ENE ‘Newhaven’ Homestead, Jul 2002, Harris 4 (NSW,
NT); 13 km ENE ‘Newhaven’ Homestead, Jul 2002, Latz
18773 & Harris (NT); Yaripilangu Range, 12 km ENE
‘Newhaven’ Homestead, Mar 1997, Latz 15097 (CANB,
DNA, NT). Queensland. Burke District: 53 km ENE
of Mt Isa, N of Breakfast Creek, Jul 1988, Harris 246
(BRI); W of Mt Isa on May Downs Station, Sep 2008,
Kleinig 6768 (AD); May Downs Station, W of Mt Isa,
Jul 2004, Booth 3563 & Kelman (BRI, CANB); 11 km
SSW of Mt Isa, Aug 1988, Harris 250 (BRI); S of Mt
Isa on the road to Dajarra, Apr 1995, Nicolle 1330 (AD);
16 km S of Mt Isa, Sybella Creek, Apr 1988, Harris 239
(BRI); South of Mt Isa on the road to Dajarra, Nov 2001,
Nicolle 4183 (AD, BRI, CANB); c. 20 km S of Mt Isa
on Dajarra Road, Sep 2008, Kleinig 6767 (AD); 20.9 km
S of Mt Isa on Dajarra road. Mar 1990, Brooker 10434

Austrobaileya 8(3): 347-355 (2011)

(BRI, CANB, MEL, NSW); 21 km S of Mt Isa on Dajarra
road. May 1994, Brooker 11888 (BRI, CANB, DNA,
NSW); 22.5 km S of Mt Isa, Aug 1973, Williams 138 &
Trapnell (BRI); 29 km S of Mt Isa on stony ridge, Sep
1951, Everist 4455 (BRI).

Distribution and habitat: The new
subspecies is of widespread but has a very
scattered distribution on arid rocky sites
from the Yaripilangu Range in the west
(about 250 km WNW of Alice Springs in
the Northern Territory), eastwards to the
Mount Isa area in north western Queensland
(Map 1). Although the Northern Territory
and Queensland populations are widely
separated geographically, it is likely that
further populations occur in the intervening
area, which is remote and botanically poorly
sampled.

In central Australia (Northern Territory),
the subspecies occurs as very scattered
populations from the Yaripilangu Range to
the Ennugan Mountains but can be locally
common or dominant, such as on the slopes
of Mt Leichhardt. It occurs on granite hills
and ranges, in open mallee vegetation.
Associated eucalypts in Central Australia
include Corymbia aparrerinja K.D.Hill
& L.A.S.Johnson, C. deserticola subsp.
mesogeotica K.D.Hill & L.A.S.Johnson,
C. eremaea (D.J.Carr& S.G.M.Carr) K.D.Hill
& L.A.S.Johnson, E. gillenii Ewart & L.Kerr,
E. normantonensis Maiden & Cambage and
E. pachyphylla F.Muell., with scattered shrubs
including Grevillea wickhamii Meisn. and
Hakea grammatophylla (F.Muell.) F.Muell.
and a groundstorey dominated by Triodia
R.Br.

In north western Queensland, the
subspecies is locally common in a small
area on the Mt Isa to Dajarra road, where
it occurs on rocky outcrops and the
intervening valleys in hilly, quite rugged
topography in gravelly red sand derived from
Sybella granite. The subspecies also occurs
as very scattered populations elsewhere on
the Mt Isa Mines Lease, where Barrs (1999)
describes the vegetation unit as ‘ Eucalyptus
melanophloia dwarf open forest’. Associated
eucalypts in the Mt Isa area include
Corymbia terminalis (F.Muell.) K.D.Hill &

Nicolle & Kleinig, Eucalyptus melanophloia

351

AD219429

D. Nicolle 5205 & M. French
Eucalyptus melanophloia subsp. nana
Western slope ol Ml Leichhardt. Northern Terr
Let 21* 46* 41* Long 132* 33' .

Malleo less than one metre tall to two metros tr
Leaves dull, grey. All parts waxy.

Dominant shrub on western slopes, ridges and
normantononsls and Grevtlba wickhamlla nd T
AD. CANB, PERTH. NT
Photos

Northern Territory. Central Australia North District.
Western slope of Ml Leichhardt.

Coord, ex GPS 2l fl 48'41*S. 132 & 33'5rE; preen I

Dominant shrub. On western slopes, ridges and fool of range in stone with
scattered Corymbut apurrerinja, Eucalyptus normantonensis and Creviliea
wlckhamii and Tnodia groundstorcy.

Malice less than one metre tall to two metres tall. Bark rough throughout, hard,
moderately fissured, dark grey to black. Leaves dull grey. All parts waxy.

D. Nicolle 5205 & M. French
Duplicates to:

Database^ in: ADHERE ghaiucn: gccc.OK)

l JuJ 2008

lllllllil

Fig. 1. Holotype of Eucalyptus melanophloia subsp. nana (Nicolle 5205 & French [AD])

352

Austrobaileya 8(3): 347-355 (2011)

Map 1 . Distribution of Eucalyptus melanophloia subsp. melanophloia (O) and subsp. nana (V).

L.A.S.Johnson, E. leucophloia subsp. euroa
L.A.S.Johnson & K.D.Hill, E. leucophylla
Domin and E. normantonensis. Scattered
Acacia shrubs are present as an under storey
(including A. hilliana Maiden and A. monticola
J.M.Black) and Triodia dominates the
groundstorey. It categorises a distinct regional
ecosystem (RE1.12.2) in Queensland that
has an ‘Of Concern’ status and an estimated
area of <10,000 ha (Accad et al. 2008; http://
www.derm.qld.gov.au/wildlife-ecosystems/
biodiversity/regional_ecosy stems/details.
php?reid=1.12.2.

Notes: Eucalyptus melanophloia subsp. nana
is distinguished from E. melanophloia subsp.
melanophloia by the consistently bushy,
multi-stemmed mallee habit and also in the
generally smaller and more crowded juvenile

leaves. The fruits also tend to be smaller and
the new subspecies more rarely produces
intermediate (and adult) leaves than the
typical subspecies.

This new subspecies is disjunct from the
nearest known locality of subsp. melanophloia
to the north-east (west of Croydon) by about
350 km. No intergrading populations are
known.

Cultivation trials of Mt Isa populations
of the new subspecies and three Queensland
provenances of subsp. melanophloia have
been undertaken since 1994 at Currency
Creek Arboretum in South Australia
(Nicolle 2000). The aim of these trials was
to determine if habit differences apparent
between the Mt Isa populations (subsp.

Nicolle & Kleinig, Eucalyptus melanophloia

353

Fig. 2. Eucalyptus melanophloia subsp. nana at Yindjirbi Range, Northern Territory (population voucher: Nicolle
5203 & French [AD, CANB, DNA]). Photo: D. Nicolle

nana) and E. melanophloia elsewhere in
Queensland (subsp. melanophloia) are
caused by environmental factors (subsp.
nana occurring in a more arid environment)
or due to genetic differentiation. These trials
indicate that subsp. nana retains both its low,
bushy, mallee habit and the generally smaller
adult leaves compared to subsp. melanophloia
(Queensland populations grown from the
Mt Moffatt, Jericho and Miles areas), which
consistently developed into taller, single¬
stemmed trees with a crown composed of
larger juvenile leaves.

Although there are no known intermediate
(‘intergrading’) populations between the two
subspecies, and although the new subspecies
is consistently distinct in habit, subspecific
rather than specific status has been applied to
the new taxon due to the allopatric distribution
of the two taxa and the overlapping distinctions
of leaf and fruit morphology. It is likely
that the two taxa would readily interbreed

if brought into geographical contact, and
the discovery of intermediate populations
remains a possibility.

Sporadic hybrids are known with
Eucalyptus normantonensis Maiden &
Cambage, of E. series Buxeales (see below).
This inter-series hybrid is conspicuous in the
field, having a crown of leaves intermediate in
morphology between the dull greyish, sessile,
broad leaves of Eucalyptus melanophloia
subsp. nana and the glossy green, petiolate,
narrow leaves of E. normantonensis (Fig. 4).

Conservation status : Although populations
are widely scattered, the subspecies is locally
common or dominant where it has been
recorded, and it is not considered to be at
risk. At Mt Leichhardt in Central Australia,
the subspecies is dominant or co-dominant on
the ridges, summit and western slopes, with
the population size likely to exceed 10,000
individuals (Fig. 3).

354

Austrobaileya 8(3): 347-355 (2011)

Fig. 3. Eucalyptus melanophloia subsp. nan a at Mt Leichhardt, Northern Territory (type locality). Photo: D. Nicolle

Etymology : From the Latin nanus (a dwarf)
referring to the consistently low bushy habit
and also in reference to the generally smaller
leaves of the new subspecies in comparison to
E. melanophloia subsp. melanophloia.

Eucalyptus melanophloia subsp. nana x E.
normantonensis hybrids

Specimens examined : Northern Territory. 4 km NE of
Mt Leichhardt, 30 km NNE of ‘Coniston’ Homestead,
Sep 2006, Duguid 22375 & Lat= (CANB, NT); SE end
of Yindjirbi Range, Coniston Station, Jul 2008, Nicolle
5204 & French (AD, CANB, DNA).

Acknowledgments

We wish to thank Ian Brooker for discussions
regarding the new taxon, staff at AD for
assisting with loans of E. melanophloia from
NT (Alice Springs), and staff at BRI and DNA
(Darwin) for allowing access to collections.
The senior author also would like to thank
Malcolm French for field assistance in Central
Australia, Max Lines of Coniston Station for
allowing and assisting with access through
Coniston Station to the Yindjirbi Range and
Mt Leichhardt, and Tony Bean for bringing
to our attention (in 2006) the occurrence of
E. melanophloia in the Northern Territory.
We also thank Paul Forster and an anonymous
referee for providing useful comments on the
manuscript.

Nicolle & Kleinig, Eucalyptus melanophloia

355

Fig. 4. Eucalyptus melanophloia subsp. nan a x E. normantonensis hybrid {Nicolle 5204 & French [AD, CANB,
DNA]). Photo: D. Nicolle

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Nicolle, D. (2000). Currency Creek Arboretum Eucalypt
Research. Vol. 1. D.Nicolle: Adelaide.

Cycas terryana P.I.Forst. (Cycadaceae), a
new species from central Queensland

Paul I. Forster

Summary

Forster, P I. (2011). Cycas terryana P.I.Forst. (Cycadaceae), a new species from central Queensland.
Austrobaileya 8(3): 356-363. A new species of Cycas from the Broadsound and Connors Ranges
in central Queensland is described, illustrated and diagnosed as C. terryana P.I.Forst. It is known
from five populations in an area of occurrence of approximately 800 km 2 and does not occur in any
conservation reserves. A conservation status of Vulnerable based on the IUCN criterion of D2 is
recommended for the species.

Key Words: Cycadaceae, Cycas , Cycas terryana , Australia flora, Queensland flora, new species,
taxonomy, identification key, conservation status

P.I.Forster, Queensland Herbarium, Department of Environment & Resource Management, Brisbane
Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: paul.forster@
derm.qld.gov.au

Introduction

The genus Cycas L. is speciose in Australia
with 28 species recognised (Hill 1998, 2004).
The species generally occur in well defined,
geographically discrete areas in few (in several
instances one) to many populations. Botanical
exploration in Queensland continues to
reveal previously undocumented populations
of Cycas ; such a case pertains to several
populations of plants in the area southwest of
St Lawrence and northwest of Marlborough in
the Broadsound and Connors Ranges.

Hill (1992) was of the opinion, albeit
without supporting evidence, that this
material represented a zone of hybridism
(“intergrading populations”) between Cycas
media R.Br. and C. ophiolitica K.D.Hill.
This hypothesis was developed further
(Hill 1996: 3), viz. “these [populations] are
morphologically intermediate between the
putative parent species and also show the
high degree of variability to be expected from
Mendelian segregation in second and later
generations”. Such introgressed populations
were considered to occur throughout the
range of the genus in Australia (Hill 1996)
and elsewhere (e.g. Hill 1994, 2008; Hill et al.
2004), in some instances between species that
he classified in different series. This ‘multiple

Accepted for publication 18 July 2011

hybridism’ hypothesis remains to be tested
using molecular markers (e.g. haplotype
networks [Schaal & Olsen 2000]) and remains
merely as a plausible supposition in some
cases. Alternative hypotheses are that these
populations are merely variations of more
broadly defined species or in some instances
represent discrete taxa.

Hill’s propensity for designating a large
number of Cycas populations as hybrids
or examples of introgression was probably
influenced by his work on eucalypts (where
this is widespread and by comparison well
documented with genetic studies) and his
mentoring by L.A.S.Johnson who together
with Lindsay Pryor was a keen supporter
of the ‘phantom’ hybrid population concept
(Pryor 1951; Pryor & Johnson 1971; Parsons &
Kirkpatrick 1972; Johnson 1976; Kirkpatrick
& Potts 1987). Eucalypts are bisexual,
often promiscuous outbreeders with non
specialist pollinators and appear to be readily
dispersed; hence, the potential for hybrids
where different taxa adjoin is high, although
recorded instances are relatively low (Griffin
etal. 1988). Populations of hybrid individuals
may lead to extinction of the parental species
via genetic pollution, although in many
cases the progeny are sterile or with reduced
viability (Levin 2000). Alternatively, these
populations may in time lead to separate
speciation events once fertility is restored

Forster, Cycas terry ana

(e.g. via allotetraploidy) and genetic change
via selection or genetic drift occurs (Levin
2000 ).

The situation in Cycas is quite dissimilar,
insomuch as the plants are dioecious, require
pollination by insects (in variations of a
mutualistic relationship) and perhaps wind in
combination (Kono & Tobe 2007), polyploidy
is unknown (Gorelick & Krystle 2011), and
distribution of the populations are limited
by dispersal (Forster 2007). As noted by
Hill (1992) it is perhaps unlikely that strong
genetic barriers to cross pollination between
different taxa exist and so the only real
barrier to hybridism/introgression is isolation
in geographic space. Populations of Cycas
and other Australian cycads are rare in the
landscape for the above reasons, quite unlike
the situation with most eucalypts. Johnson
(1959, 1961) actively promoted the concept of
widespread hybridism in cycads, especially for
Zamiaceae ( Macrozamia ) species, although
some of his examples are merely juveniles of
much larger adults (e.g. M. lucida x M. moorei ;
juveniles of what is now known as M. johnsonii
D.L. Jones & K.D.Hill) or represented discrete
taxa that he was not aware of. Speciation in
Cycas and Macrozamia is perhaps most
likely to occur via genetic drift in isolated
populations, rather than genetic selection per
se (Gorelick 2009). This is reflected in the
occurrence of similar appearing taxa, often in
a geographic replacement series of allopatric
populations. This can be interpreted as an
indication of both continuous and incomplete
speciation in a non-adaptive radiation where
the inter-relationship between environment
and morphology is unclear (Savolainen &
Forest 2005) and in cycads is probably driven
mainly by fragmentation, rather than radical
differences in habitat specialisation ( cf. Linder
2003). Either way, the tempo of speciation is
likely to be slow given the long time period
between germination and individuals reaching
maturity, together with slow turnover of
individuals within populations.

With regard to the so called Cycas media
- C. ophiolitica intermediates south of St
Lawrence, Hill (1992) was probably only aware
of two populations based on his collection
{Hill 3788 & Stanberg ) and that of Hind 2878 ,

357

or others that he examined and determined
as such. There are now five populations of
these plants known and perhaps more in what
is largely a botanically unexplored part of
central Queensland.

Neither of the putative parental species
have populations that are particularly
geographically close to the Broadsound and
Connors Ranges plants. Hill (1996) vaguely
stated that the hybrid populations were
“wherever different species grow in relatively
close proximity”. The closest population of
Cycas media is c. 9 km to the northeast (in
different habitat and geology) and the closest
population of C. ophiolitica some c. 45 km to
the east of the overall area of occupancy for the
five populations. The southern populations of
Cycas media are predominantly coastal and
occur on substrates derived from basalts, finer
grained granites or stabilised sand masses.
The northern populations of Cycas ophiolitica
are restricted to serpentinite substrates. By
comparison the Broadsound and Connors
Ranges plants occur well inland in the ranges
on substrates derived from heavily weathered
porphyritic granites.

These populations of Cycas may well
have originated from mixing of the ancestral
lineages of both C. media and C. ophiolitica
as postulated by Hill, with speciation
following the initial hybridism and subsequent
geographic isolation. However, the situation
could well be much more complex than this
and reflect multiple instances of fragmentation
and integration of populations driven by
climatic changes since the glacial cycles of
the Pleistocene. This hypothetical pattern of
change has been actively promoted as one
of the main drivers of speciation in Mexican
cycads (Vovides et al. 2003) and similar
reproductive and dispersal biology of the
Australian cycads may infer similar patterns
of allopatry (Forster 2004).

Given that the recorded populations of
these Cycas are disjunct over an area of
occurrence of at least 800 km 2 , it is likely
that the hypothesised speciation event is well
advanced with subsequent fragmentation
following an initial introgression. It is
unlikely that these populations are any less fit

358

than the core Cycas media or C. ophiolitica
populations and to all extents they appear
to exist and operate much like any other
healthy cycad populations. If of hybrid origin,
then they probably commenced as a neutral
hybrid zone with genetic compatibility and
superficially similar habitats, although such
zones are usually in close proximity to the
putative parental species (Levin 2000) which
is no longer the case here. Whatever the origin,
these populations possess character states that
enable them to be distinguished from both
Cycas media and C. ophiolitica ; hence, they
are described below as a new species. The
hypothesised species origin is also relevant
for speciation elsewhere in Cycas and other
genera of cycads, particularly Macrozamia
Miq. from Australia (Forster 2004),
Encephalartos Lehm. in Africa (Vorster et al.
2004) and Ceratozamia Brongn. and Zamia
L. in the New World (Vovides et al. 2003,
2004).

Materials and methods

The species description is based on
examination of both live plants in habitat
and herbarium collections at the Queensland
Herbarium.

Taxonomy

Cycas terryana P.I.Forst. species nova a

C. ophiolitica differt praecipue foliis valde
fragilibus (adversum non), multo parcius
carinatis et juvenilibus non glaucis, extra
colore indumenti; quum a C. media subsp.
media differentiae principals sunt folia valde
(non infirme) fragilia, maturitate indumentum
gerentia (adversum indumentum deficiens) et
valde carinata (adversum plus minusve non
vel infirme carinata), cum colore indumenti
in cataphyllis. Typus: Queensland. Port
Curtis District: Broadsound Range, 9 March
2005, P.I.Forster PIF30618 & P.J.Machin
(holo: BRI [4 sheets, $ carpological and spirit
samples]).

Arborescent cycad with stems to 3 m high
(rarely multiheaded), 12-22 cm thick. Leaves
80-190 cm long, straight and becoming wavy
towards the apex, weakly to strongly keeled
in cross-section, pale green to somewhat
glaucous green-grey above; opposing

Austrobaileya 8(3): 356-363(2011)

leaflets inserted at 20-50° to the rhachis, the
rhachis usually terminated by paired leaflets,
tomentose; petiole 17-30 cm long, 7-15 mm
diameter, dull olive grey-green, with 4-40
short teeth (pinnacanths) 1.5-2 mm long and
spaced 8-17 mm apart, sometimes spineless.
Leaflets 184-320 per leaf, 7-10 mm apart,
being evenly spaced in lower half of leaf,
then becoming more interleaved and more
strongly keeled in upper half of leaf, brittle,
margins recurved; median leaflets at 25-45°
to the rhachis, 105-230 mm long, 5-7 mm
wide, green-grey with fawn-tan bloom when
young; weakly convex in cross section,
decurrent for 5-7 mm, up to 13 mm at base
of leaf. New growth densely tomentose with
pale cream-fawn indumentum, glabrescent.
Cataphylls pungent, linear, 8-12 cm long,
densely tomentose for entire length with
pale fawn indumentum. Microsporangiate
cones elongate-ovoid, 21-28 cm long, 6.5-8
cm diameter, with dense ferruginous-brown
indumentum; microsporophylls 28-40 mm
long, fertile zone 17-28 mm long, 4-12
mm wide; apical spine antrorsely recurved,
4-10 mm long. Megasporophylls 20-33
cm long, when young with dense fawn-
tan indumentum, aging grey, eventually
glabrescent and olive-green; ovules 4 to 6;
lamina broadly triangular 50-70 mm long,
30-38 mm wide, strongly dentate with well
developed antrorse teeth 4-7 mm long, apical
spine 17—33 mm long. Seeds ovoid, 37-40
mm long, 30-35 mm diameter, sarcotesta c. 5
mm thick, weakly grey pruinose, olive green
beneath wax covering, becoming orange
when ripe. Figs. 1-5.

Additional specimens examined: Queensland.

Leichhardt District: Broadsound Range, Mar 1964,
Everist 7624 (BRI); ditto loc., Nov 2003, Forster
PIF29714 & Halford (BRI); ditto loc., Nov 2003,
Forster PIF29715 & Halford (BRI); ditto loc.. Mar
2005, Forster PIF30617 & Machin (BRI); ditto loc., Dec

2010, Mathieson MTM965 & 966 (BRI); ditto loc., Jun

2011, Forster PIF38217 & Machin (BRI). Port Curtis
District: St Lawrence towards Croydon, Aug 1990,
Hill 3788 & Stanberg (BRI, NSW); Croydon Station
- St Lawrence Road, Jun 2011, Forster PIF38218 &
Machin (BRI); 5 km W of Prospect Hill, Nov 1992,
Forster PIF12276 & Machin (BRI); near Burwood [as
‘Barwood’], Broadsound, Jun 1981, Maconochie 2757
(BRI; CANB, DNA, NY n.v.).

Forster, Cycas terryana 359

Fig. 1. Cycas terryana. Plants in habitat (population voucher: Forster PIF29714 & Halford [BRI]). Photo: P I.Forster

Distribution and habitat : Cycas terryana
is restricted to the Broadsound and Connors
Ranges northwest of Marlborough and
southwest of St Lawrence. Populations
occur on undulating low hills on stony soil
derived from porphyritic granite in woodland
dominated in various combinations by Corymbia
clarksoniana (D.J.Carr & S.G.M.Carr) K.D.Hill
& L.A.S.Johnson, C. dallachiana (Benth.)
K.D.Hill & L.A.S.Johnson, Eucalyptus crebra
F.Muell., E. drepanophylla F.Muell. ex Benth.
and Petalostigma pubescens Domin.

Notes: Cycas terryana appears to have
first been collected by Selwyn Everist in
1964; however, this herbarium specimen
has been variously identified as C. media
or an intergrade between C. media and
C. ophiolitica (determination of K.D.Hill).

The primary morphological differences
between Cycas terryana , C. media and
C. ophiolitica are summarised in Table 1 . The
leaflets and rhachis of C. terryana are very

brittle and easily snapped, as opposed to the
situation in C. media and C. ophiolitica where
they are less brittle in the former and robustly
flexible in the latter. The megasporophylls in
Cycas terryana are generally broader than in
the other species being compared here and
indumentum on the new growth, cataphylls
and megasporophylls is pale grey-fawn to
fawn-tan whereas in the others it is usually
orange-brown, only becoming greyish if
particularly weather beaten. Cycas terryana
differs from C. ophiolitica mainly in the
leaves being markedly brittle (versus not),
much less strongly keeled and not glaucous
when young, as well as the indumentum
colour; whereas from C. media subsp. media
the main differences are the leaves being
markedly brittle (versus weakly brittle), with
indumentum at maturity (versus none) and
strong keeling (versus ± flat to weakly keeled),
together with the indumentum colour on the
cataphylls.

360

Austrobaileya 8(3): 356-363(2011)

Fig. 2. Cycas terryana. Apex of mature plant showing cataphylls (population voucher: Forster PIF30617 & Machin
[BRI]). Photo: P.I.Forster

Cycas media and C. ophiolitica were
classified in different series by Hill (1996,
1998) on the basis of anatomy and morphology.
Given that species such as C. terryana exist,
there seems little point in the continued
recognition of such artificial supraspecific
taxa.

Etymology : This species is named for
Dr L. Irene Terry of the University of Utah,
U.S.A., in recognition of her ground breaking
work on cycad pollination, both in Australia
and on Guam (e.g. Terry 2001; Terry et al.
2004, 2005, 2007, 2009).

Conservation status : The species is
known from five populations with an area
of occurrence of c. 800 km and an area of
occupancy that is much less. An appropriate
status is Vulnerable based on the D2 criterion
(IUCN 2001). None of the populations occur
in conservation reserves and at least one
has been reduced to non viable numbers of
plants.

Acknowledgements

Thanks to Mike Mathieson for collecting
material and photographing the male
cones; David Halford and Peter Machin for
assistance with fieldwork and Peter Bostock
for translation of the diagnosis into Latin.

References

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Forster, Cycas terryana 361

Table 1. Comparison of character states for Cycas media, C. ophiolitica and C. terryana

Character State

C. media

C. ophiolitica

C. terryana

Mature leaf
indumentum

glabrous

tomentose below
and on rhachis

tomentose below and
on rhachis

Leaflet insertion

± flat to weakly
keeled

strongly keeled

keeled

Leaflet number

160-300

170-220

184-320

Leaflet colour
mature leaves

glossy green

glaucous grey-blue

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Leaflet texture

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flexible

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New growth
indumentum colour

pale orange-brown

mixture of grey-
white and pale
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pale grey-fawn

Cataphyll

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Megasporophyll

indumentum

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brown

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Megasporophyll
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17-30

12-30

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Seed sarcotesta
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green becoming
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Seed size: length x
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Fig. 3. Cycas terryana. Detail of megasporophylls
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Fig. 5. Cycas terry ana. Male plant with microsporangiate
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Photo: M.T.Mathieson.

A taxonomic revision of Trichosanthes L.
(Cucurbitaceae) in Australia, including one
new species from the Northern Territory

W.E. Cooper 1 & H. J. de Boer 2

Summary

Cooper, W.E. & de Boer, H.J. (2011). A taxonomic revision of Trichosanthes L. (Cucurbitaceae) in
Australia, including one new species from the Northern Territory. Austrobaileya 8(3): 364-386.
Trichosanthes is represented by six species in Australia: T. cucumerinaL. var. cucumerina, T. morrisii
W.E.Cooper sp. nov., T, odontosperma W.E.Cooper & A.J.Ford, T. pentaphylla F.Muell. ex Benth.,
T. pilosa Lour, and T. subvelutina F.Muell. ex Cogn. Trichosanthes ovigera Blume has recently been
synonymised with T. pilosa and we now include T. holtzei F.Muell. within this synonymy. All taxa are
illustrated (with the exception of T odontosperma previously illustrated in 2010), and distinguished
from other Australian species. Notes on habitat and distribution are included together with distribution
maps. Three identification keys are presented, two to the sections of Trichosanthes and one to the
species of Trichosanthes in Australia.

Key Words: Cucurbitaceae, Trichosanthes , Trichosanthes cucumerina , Trichosanthes ovigera ,
Trichosanthes pilosa , Trichosanthes odontosperma , Trichosanthes subvelutina , Trichosanthes
pentaphylla , Trichosanthes morrisii , Australia flora. Northern Territory flora, Queensland flora,
taxonomy, identification keys, new species, new combination, probracts

1 W.E. Cooper, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor
Road, Smithfield, Queensland, 4878, Australia.

2 H.J. deBoer, Department of Systematic Biology, Uppsala University, Norbyvagen 18 D, 752 36
Uppsala, Sweden. Email: hugo.deboer@ebc.uu.se

Introduction

Trichosanthes L. is a genus of approximately
100 species distributed from India, Sri
Lanka, China, Japan, SE Asia to Malesia
(including New Guinea), Australia, and
the eastern Pacific (Rugayah & de Wilde
1999). Trichosanthes is broadly represented
in Australia with one or two species from
five of the six sections: Trichosanthes L.,
Cucumeroides (Gaertn.) Kitam., Edulis
Rugayah, Aster osperma WJ.de Wilde &
Duyfjes (not in Australia), Foliobracteola
C.Y.Cheng & C.H.Yueh and Involucraria
(Ser.) Wight. Sections are distinguished using
(1) seed shape and structure, (2) colour of fruit
pulp, (3) presence or absence of probracts,
(4) size of male bracts. However, leaf shape
is variable within all species, and shape of
male bracts can vary greatly in Trichosanthes
cucumerina L., T. pentaphylla F.Muell. ex
Benth. and T. subvelutina F.Muell. ex Cogn.

The last comprehensive treatment of
Trichosanthes in Australia (Telford 1982)

Accepted for publication 14 October 2011

described six species in which T. holtzei
F.Muell. was thought to be closely allied
to T. ovigera Blume; however, T. holtzei is
no longer maintained here with the present
authors synonymising it with the widespread
and variable T. pilosa Lour. Examination of
a greater number of herbarium specimens
collected in Australia during recent years, as
well as molecular analysis (de Boer & Cooper
unpublished), have confirmed this finding.
Trichosanthes ovigera has been previously
referred to this synonymy through priority of
T. pilosa (de Wilde & Duyfjes 2008).

Telford(1982) suggestedthat Trichosanthes
pentaphylla (Fig. 1,6) is probably conspecific
with T. trifolia (L.) Merr. and the latter has
now been synonymized with T. wawrae
Cogn. (Rugayah & de Wilde 1997). However,
the two species differ sufficiently to maintain
Trichosanthes pentaphylla as distinct from
T. wawrae.

Taxa in the genus Trichosanthes can
be extremely variable throughout their
distribution with regard to leaf (Fig. 3,4),

Fig. 1 . Trichosanthes pentaphylla from the Mowbray River, showing silvery leaves with dark green veins, typical
juvenile growth of plants in the section Involucraria. Photograph: W.T. Cooper

Fig. 2. Trichosanthes subvelutina. male flowers, Terania Creek NSW. Photograph: H. Nicholson

366

bract and fruit morphology, as well as seed
surface in T. pilosa. The extremes of habitat
preferences within one species can often be
associated with morphological variation.
Trichosanthes sp. Fine Leaf (L.A. Craven
7930) had been recognised as a different taxon
by the Northern Territory Herbarium (DNA);
however, the present authors see it as part of
the widespread and variable T. cucumerina L.
var. cucumerina.

In this revision, six species of
Trichosanthes are recognised for Australia
and of those, T. odontosperma W.E.Cooper
& A.J.Ford, T. pentaphylla , T. subvelutina
and T. morrisii W.E.Cooper are endemic.
Five species are mostly tropical with one
species ( Trichosanthes subvelutina) mainly
subtropical.

Materials and methods

We examined 363 specimens from BR, BRI,
CANB, CNS, DNA, K, L, MEL, P, PERTH
and NSW. All species were observed in the
field by the first author.

Flowers of Trichosanthes are fragile and
difficult to preserve as good quality dried
specimens. They are therefore not easy to
analyze (Duyfjes & Pruespan 2004), and
have not been used as key species characters.
Even with fresh flowers, plants are mostly not
identifiable to species; however, plants having
flowers with fimbriate petal margins belong
to sections Trichosanthes or Cucumeroide s,
and those having petals with fimbriate
apices belong to sections Foliobracteola ,
Involucraria or Edulis. Leaf, probract and seed
features provide good characters (Duyfjes &
Pruespan 2004) and along with male bract
size and sepals, have been predominantly
used here as key features for dichotomous
determination keys.

Measurements of all parts were done on
dried material and specimens preserved in
70% ethanol, as well as on fresh material in
the field. Common abbreviations in the text
include N.P (National Park) and S.F./S.F.R.
(State Forest/State Forest Reserve).

The extensive bibliographic taxonomic
history of Trichosanthes sections and the
species that also occur outside of Australia has

Austrobaileya 8(3): 364—386 (2011)

been repeated several times in recent years
(e.g. Rugayah & de Wilde 1997,1999; Duyfjes
& Pruespan 2004; de Wilde & Duyfjes 2010)
and is largely omitted from this account.

Taxonomy

Trichosanthes L., Sp. PI. 2: 1008 (1753).
Type species: Trichosanthes anguina L.,fide
M.L.Green, Prop. Brit. Bot. 190 (1929) [= T.
cucumerina subsp. anguina (L.) Greb.]

Cucumeroides Gaertn., Fruct. Sem. PI. 2: 485,
t. 4 (1791). Type species: not designated.

Involucraria Ser., Mem. Soc. Phys.
Genev. 3(1): 25, t. 5. (1825). Type species:
Involucraria wallichiana Ser.

Trichosanthes in Australia: Monoecious or
mostly dioecious, trailing or climbing vines or
lianas, annual or mostly perennial, partially
or completely seasonally senescent; bark
corky, warty, flaky, fissured or lenticellate;
stems either weak and slender or robust and
somewhat woody; glabrous, pubescent or
scabrous; roots tuberous. Indumentum of
translucent multicellular trichomes, glandular
or eglandular; cystoliths often present, discoid
or rosette-shaped, drying to white or black
in herbarium specimens. Probracts present
at the nodes in many species, caducous or
persistent, may be pubescent, glands often
present. Tendrils unbranched or 2-, 3- or 4-
branched(upto 9-branched in Asia), positioned
at the node (not truly axillary). Leaves simple
or compound (digitate), alternate, petiolate,
unlobed or 3-7 (rarely 10)-lobed, margin
entire or toothed with a soft mucro or callus
terminating main veins, membranous or
leathery, scabrous, velvety or glabrous, glands
often present on the underside, pedately
veined with 3-5 main veins, intralateral veins
reticulate. Inflorescence beside petiole at the
node (not truly axillary). Flowers unisexual,
mostly nocturnal excepting T. cucumerina,
epigynous, petals 5 (rarely 4); corolla white,
cream or yellowish, actinomorphic; sepals
5 (rarely 4), free, subulate or triangular;
petals hairy, margin fimbriate; receptacle
tube elongate. Male flowers either solitary,
paired, or in racemose or rarely paniculate
inflorescences; bracts present on racemes
at pedicel base and sometimes at peduncle

367

Cooper & de Boer, Trichosanthes in Australia

base, persistent or caducous, glands may be
present; stamens 3, inserted near receptacle
tube orifice, included, free until anthesis
then becoming fused; filaments free; anthers
oblong, s-shaped, two are 2-thecous, one is
1-thecous, basifixed. Female flowers mostly
solitary or paired (one is usually much older),
sometimes in a racemose inflorescence in T.
pilosa , solitary and usually co-axillary with
male raceme in T. cucumerina ; perianth
similar to males; style long, narrow; stigma
3-5-lobed; staminodes absent; ovary inferior,
pubescent or glabrous, elliptic, ovoid, globose
or fusiform; ovules numerous, horizontal.
Fruit apepo or berry, ellipsoid, ovoid, globose
or fusiform, beaked at apex, 30-140 mm

long, mostly solitary; epicarp green with or
without longitudinal white stripes, orange or
red, glabrous or pubescent; epicarp leathery,
chartaceous or woody; mesocarp fleshy or
firm, orange, yellow or whitish, smooth or
fibrous; pulp white, yellow, orange, red or
dark green; seeds few-numerous, horizontal,
1-locular or 3-locular with the two lateral
locules empty, compressed or turgid, smooth
or sculptured, brown, cream, grey or black.
Germination is hypogeal, semi-hypogeal or
epigeal.

Etymology : The generic epithet Trichosanthes
is derived from the Greek tricho- (hairy) and
-anthus (flowered); referring to the petals,
which have fimbriate margins.

Key to sections of Trichosanthes in Australia using mostly seed and fruit features

1 Seeds 3-locular.Section Cucumeroides

1. Seeds 1-locular.2

2 Seeds surrounded by dark green pulp, adult leaves compound or simple;

juvenile leaves silvery with dark green veins.Section Involucraria

2. Seeds surrounded by white, greenish-white, yellow, orange or red pulp, all

leaves simple; juvenile leaves green only.3

3 Seeds surrounded by white or greenish-white pulp.Section Foliobracteola

3. Seeds surrounded by yellow, orange or red pulp.4

4 Seeds toothed; probract present on new growth; plants dioecious.Section Edulis

4. Seeds with an undulate margin; probract absent; plants monoecious

.Section Trichosanthes

Key to sections of Trichosanthes in Australia using leaf, probract and male bract

1 Leaves compound.Section Involucraria

1. Leaves simple.2

2 Probracts present (at least on new growth).3

2. Probracts absent.5

3 Probracts caducous.Section Edulis

3. Probracts persistent.4

4 Leaf upperside with numerous cystoliths.Section Involucraria

4. Leaf upperside without cystoliths.Section Foliobracteola

5 Plants monecious; male inflorescence bracts up to 3 mm long.... Section Trichosanthes

5. Plants dioecious; male inflorescence bracts 4-8 mm long.Section Cucumeroides

368

Austrobaileya 8(3): 364—386 (2011)

Key to species of Trichosanthes in Australia

1 Leaves compound (Qld only).T. pentaphylla

1. Leaves simple.2

2 Leaf upperside glabrous or glabrescent.3

2. Leaf upperside hairy.4

3 Probract linear or narrowly ovate, caducous; seeds toothed, surrounded

in orange or red pulp (Qld only).T. odontosperma

3. Probract obovate, elliptical or oblong, persistent; seeds entire, surrounded

in dark green pulp, (NT only).T. morrisii

4 Plants monoecious; male inflorescence bracts up to 3 mm long; seeds

with an undulate margin (NT, Qld, WA).T. cucumerina var. cucumerina

4. Plants dioecious; male inflorescence bracts 4-33 mm long; seed margin

not undulate.5

5 Male inflorescence bracts 4-8 x 0.5-3 mm; seeds tumid, t-shaped,

surrounded by yellow-orange pulp (NT, Qld, WA).T. pilosa

5. Male inflorescence bracts 15-33 x 7-23 mm; seeds flat, round and entire,

surrounded by greenish-white pulp (NSW, Qld).T. subvelutina

Trichosanthes L. section Trichosanthes

Type species: Trichosanthes anguina L. [= T.
cucumerina var. anguina (L.) Haines]

Monoecious trailing vines. Probracts absent.
Male bracts persistent (caducous in Asia).
Flowers diurnal, petals fimbriate with thread¬
like extensions along the margins. Fruit yellow
or orange; seeds 1-locular, flat, margins broad
and undulate.

Distribution : One species (two varieties)
occurring in China, India, Sri Lanka, SE
Asia, Malesia and Australia (one variety).

1. Trichosanthes cucumerina L., Sp.

PI. 2: 1008 (1753). Type: habitat in India,
“Padavalam ” in Rheede, Hort. Malab. 8: 39,
t. 15 (1688) (lecto: fide Keraudren-Aymonin
[1975: 91]).

The species comprises two varieties
as accepted here with T. cucumerina var.
cucumerina occurring in Australia. The
second variety ( T. cucumerina var. anguina
(L.) Haines) is the common snake gourd of
horticulture. Data from molecular sequencing
support the placement of the cultivated snake
gourd as a cultivated variant of T. cucumerina
(Ali & Al-Hermaid 2010).

Trichosanthes cucumerina var. cucumerina;

Haines, Bot. Bihar Orissa 3: 388 (1922).

Trichosanthes pedatifolia Miq., FI. Ind. Bat.
1: 677 (1856). Type: Indonesia: Java, s.dat.,
T.Horsfield s.n. (holo: BM28187; iso: U).

Trichosanthes reniformis Miq., FI. Ind. Bat.
1: 675 (1856). Type: Indonesia: Java, s.dat.,
T.Horsfield s.n. (holo: BM28189, 28188).

Trichosanthes ambrozii Domin, Biblioth.
Bot. 89: 631 (1929). Type: Australia: Western
Australia, between Ashburton and De Gray
Rivers, s.dat., E.Clement s.n. (syn: B, K, PR
n.v.).

Trichosanthes brevibracteata Kundu, J. Bot.
77: 10 (1939). Type: India. Punjab, Karnal, in
1885-1888, J.R.Drummond 25031 (syn: K);
India. Ahmedabad, July 1920, L.J.Sedgwick
s.n. (syn: K).

Trichosanthes pachyrrhachis Kundu, J. Bot.
77: 9 (1939). Type: Northwest India, in 1844,
M.P.Edgeworth 63 (holo: K).

Trichosanthes sp. Nitmiluk. (C.R.Mitchell
3293); Australian Plant Census (CHAH 2005);
Australian Plant Name Index, http://www.
anbg.gov.au/cgi-bin/apni. Accessed February
2010 .

Cooper & de Boer, Trichosanthes in Australia

369

Fig. 3. Trichosanthes cucumerina var. cucumerina. A. male flower. B. seeds. C. unlobed leaf underside showing
glands, male inflorescence with bracts, fruit and tendril. D. male inflorescence, buds and broadly ovate and orbicular
cupular bracts. E. leaf underside with deeply lobed margin and solitary gland. A-C from Cooper 2119 (CNS); D-E
from Fox 2513 (DNA) (scales as indicated). Del. W.T. Cooper.

Trichosanthes sp. Fine Leaf (L.A.Craven
7930), Australian Plant Census (CHAH 2005);
Kerrigan & Albrecht (2007); Australian Plant
Name Index, http://www.anbg.gov.au/cgi-bin/
apni . Accessed February 2010.

Illustrations : Yueh & Cheng (1974: pi.82);
Telford (1982: 197); Wheeler (1992: 254);
Rugayah & de Wilde (1999: 231); Hyland et
al. (2003); Duyfjes & Pruesapan (2004: 81).

Monoecious slender trailing vine to 2 m.
Stem diameter 1-2 mm, 5-ribbed, pubescent

(trichomes may be sparse); trichomes
short glandular and eglandular and mostly
interspersed with scattered long eglandular
trichomes, denser at nodes or sometimes only
at nodes; cystoliths rarely present. Probracts
absent. Tendrils 2 or 3-branched, sparse
glandular or eglandular trichomes towards
base. Leaves simple, discolorous, membranous;
petioles 8-57 mm long, sparse to dense short
glandular and eglandular trichomes, mostly
with scattered longer trichomes, rarely with a
few discoid cystoliths; lamina cordate, ovate,

370

broad-ovate or reniform, 23-145 mm long, 25-
135 mm wide; unlobed, or shallowly to deeply

3- 7-lobed, rarely 10-lobed, lobe divisions
up to c. 95% of lamina length; central lobes
oblong, elliptical, obovate or spathulate; lateral
lobes ovate, triangular, oblong, asymmetrical,
spathulate; base cordate and shortly cuneate
or truncate, sinus often shallow and wide;
apex acute, acuminate or obtuse, with a
soft mucro; margin undulate, denticulate,
dentate or serrate, 4-40 teeth or soft mucros
along each side; membranous; upperside
pubescent, hirsute or scabrous, trichomes
short eglandular and/or glandular, sometimes
restricted to main veins, and mostly with
scattered long eglandular trichomes often
restricted to near the margin, rarely with a
few scattered discoid cystoliths; underside
pubescent, hirsute or scabrous, trichomes
short glandular and/or eglandular, often
with longer scattered eglandular trichomes
usually on veins or close to margin; glands at
leaf base 1-8 per side, some scattered glands
may be present, gland diameter 0.2-0.3 mm;
rarely with a few scattered discoid cystoliths.
Male inflorescences racemose or solitary
(rarely a panicle), usually arising well after
co-axillary female; raceme 70-240 mm long,

4- 21-flowered, rachis with sparse to dense,
short glandular and eglandular trichomes as
well as long eglandular trichomes, peduncle
25-180 mm long; bracts persistent at base of
pedicels and often at base of peduncle, ovate,
broadly ovate, orbicular, triangular, narrowly
triangular, reniform, obovate, oblong,
elliptical, rhombic or linear, often cupular,
0.25-3 mm long, 0.25-2.5 mm wide, base
truncate or cuneate, apex obtuse, acute or
rarely obcordate, entire, crenate, unlobed or
with 3-5 shallow lobes, mostly orange-brown
or pinkish-brown in herbarium specimens
(linear bracts remain green), glabrous or
trichomes sparse to dense and usually much
sparser than those on the adjoining rachis,
glands absent; venation absent, obscure or
3-veined with a midrib and intramarginal
veins. Male flowers: pedicels 5-45 mm long;
receptacle tube slender-funnelform, 7-35 mm
long; trichomes glandular and eglandular,
sparse to moderately dense; sepals triangular,
entire, 0.5-2.5 mm long, 0.25-1 mm wide,
green, trichomes sparse to dense; corolla

Austrobaileya 8(3): 364-386 (2011)

diameter 8-20 mm (not including fimbrillae);
petals elliptic-ovate, 9-10 mm long (not
including numerous fimbrillae), c. 4 mm wide,
white, pubescent abaxially and adaxially;
anthers c. 2 mm long; filaments c. 0.8 mm
long. Female flowers: solitary or paired, co-
axillary with male racemes (usually emerging
before male inflorescence is fully expanded);
pedicels 3-50 mm long, trichomes glandular;
receptacle tube, slender-funnelform, 12-50
mm long, trichomes glandular and eglandular,
pubescent; sepals triangular or subulate,
entire, 1-3 mm long, 0.5-1.5 mm wide, green,
pubescent abaxially and adaxially; corolla
diameter 9-25 mm (not including fimbrillae),
petals ovate, 5-10 mm long, 2-3 mm wide,
white, trichomes glandular and sparse;
ovary 4-25 mm long, glabrous or trichomes
sparse. Fruit elongated-globular, fusiform
or globose with a beaked apex, 21-50 mm
long, 18-25 mm wide, orange or red (unripe
fruit is longitudinally streaked dark and pale
green), glabrous or glabrescent; pulp meagre,
translucent orange or reddish; mesocarp 2-2.5
mm wide; pedicel 3-9 mm long, trichomes
sparse, bract from base of male raceme usually
persistent at base; exocarp c. 0.3 mm wide;
seeds 3-12, elliptical or obovoid, 1-locular,
margin undulate with 3-5 shallow lobes on
each side, 6-10 mm long, 4-7 wide, 3-5 mm
thick, testa brown or grey. Germination type
unknown. Fig. 3.

Additional selected specimens (from 166 examined ):
Australia: Western Australia. Edge of small bay W
of Cape Ruthiers, Mar 1993, Mitchell 2989 (DNA);
c. 2 km E of Mitchell Falls, Mar 1994, Mitchell 3368
(CANB); Lacrosse Island at head of Cambridge Gulf,
NE Kimberley, Jun 1992, Kenneally 11327 (CANB);
The Dag, Mornington Wildlife Sanctuary, Apr 2005,
Legge 521 et al. (BRI); West Kimberley, Oscar Range,
Brooking Gorge, May 1988, Sands 4732 (CANB);
Dampier Archipelago, Enderby Island, Jul 1980,
Wilson 7268 (CANB); c. 135 km S of Port Hedland on
Great Northern Highway, Apr 1995, Mitchell PRP280
(CANB). Northern Territory, c. 30 km NNE of Jabiru,
Mar 1981, Craven 7930 & Whitebread (CANB); Kakadu
N.P., Mt Brockman, Mar 1995, Egan 4575 (DNA); 6 km
S Mt Gilruth, Arnhem Land, March 1984, Wightman
1345 (DNA); Deaf Adder Gorge, Feb 1977, Fox 2513
(DNA); Nitmiluk gully of Nitmiluk Gorge, Apr 2004,
Crase 1311 & Dixon (DNA); Katherine Gorge, Apr 1972,
Must 973 (DNA); 6 km NE Cape Crawford Hotel, Abner
Range, Jan 1989, Brock 484 (DNA). Queensland. Cook
District: Lankelly Creek, Mcllwraith Range, May 1995,
Hyland 15339 (CNS); Mt White, Coen, Apr 1993, Hyland
14776 (CNS); Jane Table Hill, Lakefield N.P., Mar 1993,

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Cooper & de Boer, Trichosanthes in Australia

Fell 2903 & Stanton (BRI); Metal Hills, 4 km NNE of
Chillagoe, May 2006, Wannan 4372 (NSW); Royal Arch
N.P., Chillagoe, Feb 1996, Forster PIF18614 & Ryan
(BRI); Royal Arch Tower, Chillagoe N.P., Apr 2010,
Cooper 2119 & FordJ CNS).

Distribution and habitat : Trichosanthes
cucumerina var. cucumerina is widely
distributed from India, Southern China and
Southeast Asia through Malesia to Australia.
Within Australia it is widespread in Western
Australia from the northern Kimberley south
to the Port Hedland area, also in the Northern
Territory from Arnhem Land to the Abner
Range near McArthur, and in Queensland
from Cape York to Chillagoe (Map 1). In
Queensland it is more restricted with a few
isolated collections from Cape York Peninsula
and Chillagoe-Mungana Caves N.P, and
one collection from Black Mountain near
Cooktown within the Wet Tropics bioregion.
It has a strong association with sandstone
habitats but it also occurs in areas of granite
and limestone. It is a climber on rocks, shrubs
and trees in open forest, woodland, shrubland
(often on sandstone escarpments), monsoon
forest, vine thickets, Allosyncarpia forest,
herblands and grasslands.

Phenology : Flowers have been recorded in
January, February, March, April, May, June,
July and October; fruit has been recorded in
January, February, March, April, May, June,
August and October.

Notes : Trichosanthes cucumerina var.
cucumerina is a highly variable species with
leaves varying from unlobed to deeply lobed.
However, all plants have male bracts up to 3 mm
long, flowers that are open during the day, and
seeds with undulate margins. It differs from
all other Australian Trichosanthes by having
flowers that open during daylight, and very
small male bracts. Male inflorescence bracts
are usually persistent through to ripe fruit in
Australian populations, whereas in Malesia
and China the bracts are absent, subpersistent
or caducous (Backer & Bakhuizen van den
Brink 1963; Rugayah & de Wilde 1997; de
Wilde & Duyfjes 2004, 2010; Duyfjes &
Pruesapan 2004; Lu et al. 2011). Leaf glands
are usually present in Australian populations
whereas in Malesia leaf glands are absent or
few (de Wilde & Duyfjes 2010). The leaves

have a pungent smell like those of Momordica
charantia L. mixed with peanut butter.

Etymology : After its similarity to the related
genus Cucumis.

Trichosanthes section Cucumeroides

(Gaertn.) Kitam., J. Jap. Bot. 19: 35 (1943);
Cucumeroides Gaertn., Fruct. Sem. PI. 2: 485,
t. 4, f. 4. (1791). Type species: T. cucumeroides
(Ser.) Maxim. [= T. pilosa Lour.]

Dioecious (sometimes monoecious) vines;
probract absent; male bracts persistent
(caducous or persistent in Thailand, Duyfjes &
Pruesapan 2004), relatively short and mostly
narrow; flowers nocturnal, petals fimbriate
with thread-like margins; fruit pulp yellow
or orange (also whitish or pinkish in Asia,
de Wilde & Duyfjes 2004); seeds 3-locular,
t-shaped, quadrangular and tumid.

Distribution : Approximately 14 species in
China, India, SE Asia, Malesia and Australia
(one species).

2. Trichosanthes pilosa Lour., FI. Cochinch.
1: 588 (1790). Type: Vietnam (holo: lost);
Vietnam. Tu Phap, s.dat.. Bon 4019 (neo: P,
fide de Wilde & Duyfjes [2008: 270]).

Trichosanthes ovigera Blume, Bijdr. FI. Ned.
Ind. 15: 934 (1826). Type: Indonesia: Java. Mt
Salak, s.dat., C. Blume s.n. (holo: LI30442;
iso: LI 30439).

Trichosanthes chinensis Ser. in DC., Prodr.
3: 308 (1828). Type: tab. 13 in Cattley, Ic. PI.
China (1821).

Bryonia cucumeroides Ser. in DC., Prodr. 3:
308 (1828); Trichosanthes cucumeroides (Ser.)
Maxim, in Franch. & Sav., Enum. PI. Jap.
1: 172 (1873). Type: Japan, Cucumeroides
Thunb. ex. Gaertn., Fruct. 2: 485 (1791).

Trichosanthes horsfieldii Miq., FI. Ned. Ind. 1:
677 (1856). Type: Indonesia: Java. Priangan,
s.dat., T.Horsfield s.n. (holo: BM; iso: K, U).

Trichosanthes hearnii F.Muell. ex
Benth., FI. Austral. 3: 315 (1867). Type:
Australia: Queensland. Rockingham Bay,
s.dat., J.Dallachy s.n. (syn: K; isosyn:
MEL100085).

Trichosanthes himalensis C.B.Clarke in
Hook.f., FI. Brit. India 2: 608. (1879). Type:

372

Austrobaileya 8 ( 3 ): 364—386 ( 2011 )

Fig. 4. Trichosanthespilosa. A. seed: lateral and adaxial view. B. seed: adaxial view. C. male inflorescence with bracts
and buds. D. lobed leaf underside showing glands near base. E. unlobed leaf upperside, male inflorescence and tendril.

A & E from Gray 7477 (CNS); B from Cooper 2120 (CNS):
2064 (CNS) (scales as indicated). Del. W.T. Cooper.

India: Sikkim. Yoksun to the plains, ann. coll,
ign., J.D. Hooker s.n. (syn: K); India. Sikkim,
s.dat., Clarke s.n. (syn: K).

Trichosanthes dicoelospermum C.B.Clarke in
Hook.f.,/ 7 /. Brit.Ind. 2:609(1879); T. cucumeroides

C from McDonald 8839 & Dennis (CNS); D from Cooper

var. dicoelosperma (C.B.Clarke) S.K.Chen, Bull.
Bot. Res. 5: 118 (1985). Type: India: Sikkim.
Khasia, s.dat., J.D.Hooker s.n. (syn: K); loc. cit.,
s.dat., J.D.Hooker & T. Thomson s.n. (syn: K,
UPS).

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Cooper & de Boer, Trichosanthes in Australia

Trichosanthes holtzei F.Muell. Aust. J. Pharm.
1: 447 (1886). Type: Australia: Northern
Territory, near Port Darwin, s.dat., M.Holtze
432 (holo: MEL100084), syn. nov.

Trichosanthes cavaleriei Lev., FI. Kouy-
Tcheou 123 (1914). Type: China: Guizhou
Province: near Guiyang, Bodinier 2306,
Cavalerie 1032,1816 (syn: all E).

Trichosanthes vanoverberghii Merr., Philipp.
J. Sci., C 9: 458 (1915). Type: Philippines.
Luzon Province: Bontoc subprovince, Bauco,
s.dat., M.Vanoverbergh 3662 bis (iso: K).

Trichosanthes baviensis Gagnep., Bull.
Mus. Hist. Nat. Paris 24: 379 (1918). Type:
Vietnam. Tu Phap, s.dat.. Bon [as Balansa ]
4016 (holo: P).

Trichosanthes pierrei Gagnep., Bull. Mus.
Hist. Nat. Paris 28:380 (1918). Type: Vietnam.
Baria, Mt Lu Dinh, s.dat., Pierre 4491 (holo:

P).

Trichosanthes hainanensis Hayata, Icon.
PI. Formosan. 10: 8 (1921); T. cucumeroides
var. hainanensis (Hayata) S.K.Chen, Bull.
Bot. Res. 5: 117 (1985). Type: China. Hainan
Province: s.loc., s.dat., Katsumada s.n. (holo:
TI).

Trichosanthes formosana Hayata, Icon. PI.
Formosan. 10: 7 (1921). Type: Taiwan. Taipei,
Urai, s.dat., Matsuda 267 (holo: TI).

Trichosanthes matsudai Hayata, Icon. PI.
Formosan. 10: 10 (1921). Type: Taiwan.
Qishan, Kosenpo, s.dat., Matsuda s.n. (holo:
TI).

Trichosanthes boninensis Nakai ex Tuyama,
Bot. Mag. Tokyo 50: 133 (1930). Type: Japan.
Bonin Island, Titizima, s.dat., Nakai s.n.
(holo: TI).

Trichosanthes chingiana Hand.-Mazz.,
Sinensia 7: 621 (1936). Type: China: Guangxi
Province: Yema Shan, s.dat., R.C.Ching 7113
(holo: W).

Trichosanthes rostrata Kitam.,Acta Phytotax.
Geobot. 5: 210 (1936). Type: Japan. Ryukyu
Islands. Miyagi Island, s.dat., Koidzumi s.n.
(holo: KYO n.v., iso: US36699).

Trichosanthes cucumeroides var. stenocarpa
Honda, Bot. Mag. Tokyo 54: 223 (1941). Type:
Japan. Musasi, Urawa, s.dat., K.Emori s.n.
(holo: TI).

Trichosanthes okamotoi Kitam., J. Jap. Bot.
19: 40 (1943). Type: Taiwan. Gaoxiong,
‘Benchi, Chushinron’, s.dat., Okamoto s.n.
(holo: KYO).

Trichosanthes mafuluensis Merr. & L. M. Perry,
J. Arnold Arbor. 30: 58 (1949). Type: Papua
New Guinea: Central Province. Mafulu,
1933, L.J.Brass 5257 (holo: A; iso: BO, BRI).

Trichosanthes ascendens C.Y.Cheng &
C.H.Yueh, Act. Phytotax. Sin. 18: 340 (1980).
Type: China: Guangxi Province: He Xian,
s.dat., C.T. Li 604120 (holo: IBX).

Trichosanthes trichocarpa C.Y.Wu ex
C.Y.Cheng & C.H.Yueh, Acta Phytotax.
Sin. 18: 340 (1980). Type: China. Yunnan
Province: Baoshan Xian, s.dat., T.T. Yu 17861
(holo: KUN; iso: A).

Illustrations: Keraudren-Aymonin (1975: 87)
as T. ovigera, Telford (1982:197) as T. ovigera,
Brennan (1986: 26) as T. cucumerina. ; Jones &
Gray (1988: 354) as T. holtzei, Wheeler (1992:
254) as T. ovigera, Rugayah & De Wilde
(1999: 231) as T. ovigera, Hyland et al. (2003)
as T. ovigera, Duyfjes & Pruesapan (2004: 81)
as T. ovigera. Cooper & Cooper (2004: 144)
as T. ovigera. Cooper & Cooper (2004: 144)
as T. holtzei.

Dioecious or monoecious trailing vine to 6

m, perennial, partly or completely seasonally
senescent. Stem diameter 1-3 mm, 5-
ribbed, glabrous or glabrescent, trichomes
glandular and/or eglandular, scattered longer
eglandular trichomes may be present; nodes
usually slightly swollen, discoid cystoliths
may be present at nodes. Probracts absent.
Tendrils unbranched or 2- or 3-branched.
Leaves simple, discolorous, membranous
or chartaceous; petioles 17-85 mm long,
eglandular trichomes present, interspersed
with sparse glandular trichomes, rarely with
a few longer trichomes, discoid cystoliths
may be present; lamina cordate, unlobed or
shallowly to deeply 3-lobed, 70-200 mm long,
50-155 mm wide, base cordate, usually shortly
cuneate, sinus deep and narrow (rarely wide),

374

apex acute, acuminate, obtuse or retuse, with
a soft mucro, margin remotely denticulate,
undulate or crenate (mostly almost entire
except for a small mucro terminating each
vein), 16-28 mucros on each side; upperside
pubescent, hirsute or scabrous, sparsely to
densely clothed in short eglandular trichomes,
rarely with some glandular trichomes near
base, sometimes interspersed with longer
trichomes which are often only present near
margin; underside pubescent, hirsute or
scabrous, sparsely to densely covered in short
eglandular or glandular trichomes (usually
denser on underside), often with a few longer
trichomes on main veins especially towards
margin; 1-5 glands at leaf base on each side,
rarely with scattered glands, gland diameter
0.2-0.9 mm; discoid cystoliths may be present
on main veins. Male inflorescence racemose
or sometimes co-axillary with a solitary
flower; raceme up to 200 mm long and up to 11-
flowered, rachis densely covered in glandular
trichomes, peduncle 15-75 mm long; bracts
persistent, lanceolate, ovate, rhomboid, linear,
elliptical or obovate, 4-8 mm long, 0.5-3
mm wide, base truncate or cuneate, apex
acute or rarely obtuse, margin entire or 3-7-
toothed, glabrous or glabrescent, trichomes
glandular and eglandular, glands absent,
discoid cystoliths may be present, venation
pinnate and reticulate. Male flower: pedicels
5-35 mm long; receptacle tube salverform or
slender-funnelform, 23-24 mm long, clothed
in glandular and eglandular trichomes; sepals
triangular, subulate or lanceolate, entire or 3-
toothed, 4-7 mm long, 1-2 mm wide, green,
clothed in glandular and eglandular trichomes;
corolla diameter 12-20 mm; petals ovate, 6-8
mm long (not including fimbrillae), white,
cream, yellow or greenish-yellow, pubescent.
Female flowers in racemose inflorescence or
solitary (rarely intermixed on a male raceme)
on a pedicel 10-30 mm long; racemes 30-
70 mm long, 5-10 flowered; bracts present
on racemes and similar to those on male
inflorescences, trichomes glandular and
eglandular. Female flower: pedicels 12-16
mm long for solitary flowers, c. 5 mm long for
flowers in racemose inflorescence, trichomes
glandular and eglandular; receptacle tube
salverform or slender-funnelform, 18-50 mm

Austrobaileya 8(3): 364-386 (2011)

long (including ovary), trichomes glandular
and eglandular; sepals triangular or subulate,
often recurved, 2-9 mm long, 0.5-3.5
mm wide, green, trichomes glandular and
eglandular; corolla diameter (not including
fimbrillae) 15-17 mm; petals ovate, 8-18 mm
long, 4-9 mm wide (not including fimbrillae),
white or cream, clothed in glandular and
eglandular trichomes; fimbrillae white,
yellow or greenish-yellow; ovary 10-16 mm
long, trichomes glandular and eglandular.
Fruit obovoid or oblong-globose, apex acute
or obtuse with a short beak, 10-ribbed (ribs
sometimes subtle), 31-62 mm long, 31-35
mm wide, orange or red, glabrous or a few
glandular trichomes near base; mesocarp 2-
2.5 mm wide, orange; pedicel 7-10 mm long;
seeds 10-21, t-shaped, subquadrangular,
tumid, 3-locular, central cell warty to smooth
and containing the embryo, lateral cells cone-
shaped and empty, 5-10 mm long, testa brown
or blackish, suspended in yellow-orange pulp.
Germination is hypogeal or semi-hypogeal,
cotyledons straight. Fig. 4.

Additional selected specimens {from 85 examined ):
Western Australia. Mitchell Plateau, N Kimberley, Feb
1979, Beard 8436 (PERTH). Northern Territory. Gurig
N.P, Apr 2006, Brennan 6919 (DNA); Murgenella,
Wunyu Beach, Mar 1987, Russell-Smith 1967 & Lucas
(DNA); 5 km E mouth of Peter John River, NE Arnhem
Land, Feb 1988, Russell-Smith 4768 & Lucas (DNA);
Rocky Bay, Yirrkala, Mar 1988, Russell-Smith 5168 &
Lucas (BRI); 1 km SE of Angurugu, Groote Eylandt,
Mar 1988, Russell-Smith 5135 & Lucas (CANB); 5 km
SW Cutta Cutta Caves, Feb 1989, Russell-Smith 7157 &
Lucas (DNA); Van Dieman Gulf at base of Cape Hotham
Peninsula, Mar 1993, Cowie 3291 (DNA); Gunn Point,
Apr 1984, Dunlop & Wightman 6678 (DNA); Wangi
Falls, Litchfield N.P., Dec 2009, Cooper 2100 & Morris
(CNS). Queensland. Cook District: Old Mapoon Road,
Jun 2000, Hyland 16381 (CNS); Iron Range N.P., Mar
1994, Fell 4118 & Stanton (BRI); Lake Patricia, Weipa,
Mar 1989, O’Reilly 262 (BRI); Turrell Hill, Macrossan
Range, Silver Plains, Jun 1998, Forster PIF23085
et al. (BRI); Brooklyn, May 2010, Cooper 2120 &
Russell (CNS); Barron Gorge, Aug 2001, Cooper 1569
& Cooper (CNS); Eight Mile Mt, Mar 1999, Gray 7477
(CNS); Burke Developmental Road, Royal Arch Tower,
Chillagoe N.P, Feb 2010, Cooper 2102 & Ford (CNS);
Barron Gorge N.P, Apr 2009, Cooper 2064 (CNS);
Kennedy Highway, 300 m N of Herberton turnoff. May
2004, Cooper 1840 & Cooper (BRI); N.P.R. 16, Forty
Mile Scrub, old road. Mar 1999, Ford 2193 (CNS); Wind
Tunnel Lava Tubes, Undara N.P, Mar 2010, McDonald
8839 & Dennis (CNS). South Kennedy District: Carlisle
Island, Sep 1986, Batianoff5038 (BRI).

375

Cooper & de Boer, Trichosanthes in Australia

Distribution and habitat : Trichosanthes
pilosa is widely distributed from India through
Southern China to Japan, Southeast Asia
through Malesia to Australia and the eastern
Pacific. Within Australia it is widespread in
the Northern Territory and in Queensland it
occurs from Cape York south to Carlisle Island
(northeast of Mackay) and west to Chillagoe
(Map 2). There is one specimen from the
Mitchell Plateau in the northern Kimberly area
of Western Australia. It occurs in a variety of
habitats from wet tropical rainforest, monsoon
forest and littoral rainforest to deciduous vine
thickets at an altitudinal range from near sea
level to 1000 m.

Phenology: Flowers have been recorded in
February, March, April and June; fruit has
been recorded in February, March, April,
May, June and August.

Notes : Trichosanthes holtzei F.Muell. was
described as being reminiscent of T. anguina
(synonym of T. cucumerina var. anguina)
by Mueller (1886). His description is of a
specimen with staminate and pistillate flowers.
However, pistillate flowers are not evident on
the type specimen today, and even though
pistillate flowers may have been present,
this feature is not sufficient to distinguish
it, as Trichosanthes pilosa can at times be
monoecious. The inflorescence bracts and
leaves on the type specimen are a good match
for Trichosanthes pilosa , and T. holtzei is here
synonymised with T. pilosa.

A second variety of Trichosanthes pilosa
was described by de Wilde & Duyfjes (2008)
restricted to Thailand. Molecular analyses to
date do not support the recognition of varieties
in this species (de Boer, unpublished). Only
the nominative variety occurs in Australia.

Etymology : The specific epithet pilosa is
derived from the Greek pilos (anything
made of felt), presumably referring to the
indumentum.

Trichosanthes section Edulis Rugayah,
Reinwardtia 11: 232 (1999). Type species:
T. edulis Rugayah

Dioecious trailing vines or lianas; probract
present, caducous; flowers nocturnal, petal
margin fimbriate in upper half; fruit pulp

red; seeds numerous, 1-locular, flat or
quadrangular, toothed or notched.

Distribution : Nine species (New Guinea eight
endemic, Australia one endemic).

3. Trichosanthes odontosperma W.E. Cooper
& A.J.Ford, Austrobaileya 8: 126 (2010).
Type: Australia: Queensland. Cook District:
Topaz, Westcott Road, 15 April 2009,
W.Cooper 2065 (holo: CNS; iso: BRI, CANB,
DNA, L, MO, NSW, UPS).

Illustrations: Cooper & Cooper (1994: 291),
as Trichosanthes sp. (Mt Lewis); Cooper &
Cooper (2004: 145), as Trichosanthes sp. (Mt
Lewis); Hyland et al. (2003), as Trichosanthes
sp. (Mt Lewis BG 167); Jones & Gray (1988:
354 & back cover), as Trichosanthes sp.;
Williams (1987: 307), as Trichosanthes sp.;
Cooper & Ford (2010: 128-129).

Dioecious trailing vine or liana to mid¬
canopy, perennial, partly or completely
seasonally senescent. Stem diameter to 3 cm,
young branchlets 5-angular, glabrescent with
minute trichomes clustered at nodes; bark
fissured and corky on older growth; nodes
often markedly swollen. Probracts caducous,
linear or narrowly ovate, minutely lobed or
with a few teeth, 3-13 mm long, 1-3 mm wide,
glandular, glabrous or glabrescent. Tendrils
unbranched or 2- or 3-branched. Leaves
simple, discolorous, coriaceous; petioles
20-95 mm long; lamina ovate, cordate or
triangular, unlobed or rarely 3-lobed, 50-190
mm long, 41-160 mm wide, base cordate or
rarely hastate, sinus mostly narrow and deep,
apex acuminate to acute, with or without a soft
mucro; margin denticulate with 13-27 teeth
per side; upperside smooth, glabrous or with
sparse minute translucent trichomes on main
veins; numerous small translucent and sunken
multicellular (rosette-shaped) cystoliths,
which in most dried specimens become black;
underside with sparse translucent minute
trichomes on main veins, 1-16 circular and
flat glands on each side of the leaf base. Male
inflorescences with flowers mostly solitary,
rarely in a fascicle of two flowers or a raceme
beside a solitary flower, peduncle 35-110 mm
long; bracts at the base of racemose flowers
narrowly ovate, glabrous, 5-17 mm long,
1-3 mm wide. Male flowers: 45-90 mm

376

diameter, pedicel 43-83 mm long; receptacle
tube salverform, 45-90 mm long, green or
creamy-green with a narrow bright yellow
centre, glabrous or glabrescent abaxially;
sepals usually 5 (rarely 4), triangular, entire
or with 1-3 teeth, 8-20 mm long, 2-4 mm
wide, green, glabrous or glabrescent; petals
5 (rarely 4), obdeltoid, length including
fimbrillae 27-40 mm, 24-40 mm wide, white,
both surfaces villous becoming glabrescent
towards apex. Female inflorescences a
solitary flower on pedicels 31-50 mm long,
bracts absent. Female flowers: 60-73 mm
long, 55-65 mm diameter; receptacle tube
salverform, 39-55 mm long, green or creamy-
green with a narrow bright yellow centre,
adaxial surface of tube yellow, glabrous or
glabrescent abaxially; sepals 5, triangular,
entire, glabrous or glabrescent, 5-12 mm
long, 1-2 mm wide, green; petals 5, obdeltoid,
28-31 mm long, white, both surfaces villous;
ovary 14-24 mm long, glabrous and 10-
ribbed externally. Fruit ovoid or ellipsoid,
apex beaked, 90-140 mm long, 60-90 mm
diameter, glabrous, longitudinally 10-ribbed,
orange to red; mesocarp firm, 10-13 mm
thick, yellow-orange; pedicel 25-40 mm long
(or more, based upon flowering specimens), 4-
10 mm wide; seeds numerous, quadrangular,
1-celled, 2-horned at one end, 4-lobed at the
other end, both sides with 2 rows of 2-10
teeth, 12-18 mm long, 6.3-77 mm wide, 3-4
mm thick, testa brown to blackish, suspended
in orange or red pulp. Germination epigeal.

Distribution and habitat : Trichosanthes
odontosperma is endemic to the rainforests
of the Wet Tropics bioregion in north-east
Queensland. It occurs from the Windsor
Tableland area, west of Cape Tribulation, to
the Cardwell Range, west of Tully (detailed
notes were given previously in Cooper & Ford
[ 2010 ]).

Phenology : Flowers have been recorded in all
months; fruits have been recorded in January,
March, April, June, July, August, October and
November.

Etymology : The specific epithet is derived
from the Greek, odonto (tooth) and -sperma
(seed) and refers to the toothed seeds,
distinguishing it from all other Australian
Trichosanthes.

Austrobaileya 8(3): 364-386 (2011)

Trichosanthes section Foliobracteola

C.H.Yueh & C.Y.Cheng, Act. Phytotax. Sin.
12: 427 (1974). Type species: T. kirilowii
Maxim.

Dioecious vines or lianas; probracts present;
male bracts large; flowers nocturnal, petal
margin fimbriate in upper half; fruit pulp
whitish, cream or greenish-white; seeds 1-
locular, flat, margined.

Distribution : 12 species in China, India, SE
Asia, Malesia and Australia (one species).

4. Trichosanthes subvelutina F.Muell. ex
Cogn., Monogr. Phan. [A.DC. & C.DC.] 3:
366 (1881). Type: Australia: Queensland.
Moreton District: Moreton Bay, Archers
Brush, [November 1843], L.Leichhardt 4
(lecto [here designated]: P731577, photo!);
isolecto: P731576, photo!; MEL100124,
photo!; MEL100125, photo!).

Illustrations : Telford (1982: 197); Harden
(1990: 448); Nicholson & Nicholson (2004:
65); Harden et al. (2007: cover, 95, 96).

Dioecious trailing vines or lianas to 8

m, sometimes extending into the canopy,
perennial, seasonally senescent. Stemdiameter
to 15 mm, ribbed with vertical lenticels, bark
shallowly corky on older stems; young stems
glabrous and 5-angular, diameter c. 3 mm,
clothed in rusty trichomes, discoid cystoliths
may be present, nodes swollen. Probracts
persistent, ovate, quadrate or triangular,
entire or apex 3-toothed, 1-5 mm long, 1-4
mm wide, sparsely to densely pubescent, often
glandular. Tendrils 2-7-branched. Leaves
simple, discolorous, membranous; petioles
22-110 mm long; trichomes translucent
white, fawn or pale rusty eglandular; discoid
cystoliths may be present; lamina cordate or
broadly ovate, 3-5-lobed (rarely 7-lobed),
lobes shallow to deep, 85-260 mm long, 60-
220 mm wide; base cordate, shortly cuneate
or truncate, sinus deep and narrow or wide
and shallow; apex acute, acuminate or obtuse,
usually with a soft mucro; margin denticulate,
undulate or almost entire except for a short soft
mucro terminating each vein, 22-37 mucros
per side; upperside pubescent (sometimes
scabrous) with translucent white or fawn
eglandular trichomes, may be interspersed

Cooper & de Boer, Trichosanthes in Australia

377

Fig. 5.. Trichosanthes subvelutina. A. leaf, probract and tendril. B. rhomboid bract. C. broad-elliptic bract. D. seeds:
left, lateral view; right, adaxial view. A & D from Nicholson 3110 (BRI); B from Boorman s.n. (NSW146961); C from
Telford 9723 (CANB) (scales as indicated). Del. W.T. Cooper.

with longer trichomes; underside pubescent
with translucent white or fawn glandular and
eglandular trichomes, glands at leaf base up
to 4 per side or absent, gland diameter 0.4-0.5
mm; cystoliths absent. Male inflorescences
racemose, paired, solitary or a solitary flower
co-axillary with a raceme, racemes c. 250 mm
long with up to 12 flowers; peduncle 75-165
mm long, pubescent; bracts persistent, broad-
elliptic, rhomboid or obovate, 15-33 mm
long, 7-23 mm wide, apex acute or obtuse,
base truncate or cuneate, densely pubescent,
some trichomes glandular, margin denticulate
or almost entire except for soft mucros,
palmately veined, reticulate, glands sparse
or absent, gland diameter 0.3-0.5 mm. Male
flowers: pedicels 35-155 mm long; receptacle
tube funnelform or slightly urceolate, 31-
45 mm long, green with fawn pubescence
abaxially; sepals triangular, entire, 8-11

mm long, 2-3 mm wide, green, pubescent;
corolla diameter 30-70 mm; petals obovate
or obdeltoid, 18-21 mm long (not including
fimbrillae), white, pubescent on both sides.
Female inflorescences with flowers solitary
or paired (one usually much older) on pedicels
18-50 mm long, pubescent. Female flowers:
receptacle tube tubular, 22-51 mm long,
pubescent, trichomes fawn or rusty; sepals
triangular, entire, 5-11 mm long, 1-3 mm
wide, green, pubescent; corolla diameter
(not including fimbrillae) 40-67 mm; petals
obovate, c. 22 mm long, white, trichomes
mostly on veins adaxially, glabrous abaxially;
ovary 18-20 mm long, pubescent. Fruit
ellipsoid, apex beaked, 75-160 mm long,
60-100 mm wide, green with paler irregular
stripes, sparsely pubescent; mesocarp firm,
c. 5 mm wide, whitish; pedicel 40-92 mm
long, diameter 3.5-6 mm; seeds numerous

378

(about 100), broad-ovate or broad-elliptical,
9-12 mm long, 6-9 mm wide, testa cream-
coloured, marginate, flesh around seeds
white or greenish-white. Germination type
unknown. Silky Cucumber. Figs. 2 & 5.

Additional selected specimens (from 35 examined ):
Queensland. North Kennedy District: Gloucester
Island, Apr 1994, Batianoff 9404129 & Figg (BRI).
Wide Bay District Veteran S.F. [now Gympie N.P.],
c. 9 km NNE of Gympie, May 1999, Bean 14829
(BRI); Booloumba Creek area, Feb 2001, Wright s.n.
(BRI [AQ552068]); Upper Kadanga S.F., 9 km NE
of Gallangowan, Oct 2008, Forster PIF34404 et al.
(BRI); Western Road, Montville, Apr 1959, Gowlett
s.n. (BRI [AQ310073]). Moreton District: Dunumbar
Reserve, London Creek, off McDonalds Road, 2 km
NNE of Peachester, Jul 2006, Forster PIF31757 &
Smyrell (BRI); Geebung, Brisbane, Jan 1977, Gray
s.n. (BRI [AQ310082]); Daisy Hill S.F., Logan City,
Dec 1992, Thompson LOG75 (BRI); O’Possum Creek,
Springfield near Ipswich, Apr 1994, Bird s.n. (BRI
[AQ627235]); Springbrook Plateau, Purlingbrook Falls,
Apr 1984, Telford 9723 (CANB); Currumbin Creek
Road to Tomewin Gap, Jan 1984, Telford 9103 (CANB);
Mt Nathan, Mudgeeraba, May 1963, Morgan s.n. (BRI
[AQ310080]); Lyrebird Ridge Road, Springbrook, Feb
2004, Edginton & Halford s.n. (BRI [AQ763103]); Binna
Burra, Feb 2004, Fechner et al. s.n. (BRI [AQ578156]).
New South Wales. Billynudgel, Feb 1913, Boorman s.n.
(NSW146961); Whian Whian S.F., N of Lismore, Apr
1939, Jouberts.n. (NSW146959); 10.2 km along Terania
Creek Road from The Channon, Apr 2009, Nicholson
3110 (BRI); Victoria Park near Rous, S of Alstonville,
Nov 1964, Williams K22 (NSW).

Distribution and habitat : Trichosanthes
subvelutina is endemic to eastern Australia.
Apart from a highly disjunct population
on Gloucester Island east of Bowen, it is
restricted to southeast Queensland (extant
northern limit at Gympie N.P.) south to Rous
near Alstonville in northern New South Wales
(Map 2). It inhabits open or closed canopy
Eucalyptus forest, disturbed rainforest,
rainforest margins and Araucarian vineforest
at altitudes from near sea level to 600 m.

Phenology : Flowers have been recorded
in January, February, March, April, June,
October and November; fruit has been
recorded in January, February, March, April
and May.

Typification : A diverse range of specimens
were cited in the original description of this
species. These were widely distributed by
Mueller to herbaria in Europe with some
(but not all) having what are presumably

Austrobaileya 8(3): 364-386 (2011)

duplicates retained at MEL. We have selected
as lectotype of this name, a fertile collection
that is labelled as being from “Archers Brush”
and collected by Leichhardt.

Other residual syntypes are as follows
- Australia: Queensland. Moreton District:
Three Mile Scrub [Ashgrove Avenue,
Ashgrove], Burrgan, Moreton Bay, July
1843, F. Mueller s.n. (BR6606213, photo!;
MEL100126, photo!); Moreton Bay, s.dat.,
L.Leichhardt 8 (P731578, photo!); Moreton
Bay, Archers Brush, November [71843],
L.Leichhardt 26 (P731579, photo!); Moreton
Bay, Archers Brush, November-December
[71843], L. Leichhardt s.n. (P731580, photo!);
Moreton Bay, s.dat., F. Mueller s.n. (P731573,
photo!); Brisbane River, Moreton Bay, s.dat.,
F. Mueller s.n. (P731575, photo!; MEL100128,
photo!); New South Wales. Clarence River,
1861, C.Moore s.n., (P731574, photo!).

Etymology : The specific epithet is derived
from the Latin sub- (almost or somewhat) and
velutinus (velvety), probably referring to the
indumentum.

Trichosanthes section Involucraria (Ser.)
Wight, Madras J. Lit. Sci. 12: 52 (1840);
Involucraria Ser., Mem. Soc. Phys. Geneve
3: 27, t. 5 (1827). Type species: Involucraria
wallichiana Ser. [= Trichosanthes wallichiana
(Ser.) Wight]

Dioecious vines or lianas; probracts persistent;
leaves simple or compound; juvenile leaves
silvery with dark green veins; male bracts
medium or large, glandular; petal margin
fimbriate in upper half; fruit red, pulp dark
green; seeds 1-locular, flat or slightly swollen,
with a narrow margin or not margined.

Distribution : Approximately 50 species in
China, India, SE Asia, Malesia (including
New Guinea) and Australia (two species).

379

Cooper & de Boer, Trichosanthes in Australia

5. Trichosanthes pentaphylla F.Muell. ex
Benth., FI. Austral. 3: 314 (1867). Type:
Australia: Queensland. North Kennedy
District: “Mt Grame” [Mt Graham, west
of Rockingham Bay], June 1864, [probably
J.Dallachy s.n. for F.Mueller] (lecto [here
designated]: MEL100120, photo!).

Illustrations : McCubbin(1971: cover, 58-59);
Jones & Gray (1988: 354); Cooper & Cooper
(1994: cover, 290); Hyland et al. (2003);
Cooper & Cooper (2004: 145).

Dioecious vine or liana to canopy, perennial,
seasonally senescent. Stem diameter to 40 mm,
bark on older stems slightly flaky, fissured,
lenticellate, not ribbed, adventitious roots
at nodes when trailing; young branchlets 7-
ribbed, diameter 4-6 mm, glabrescent or some
trichomes clustered at nodes; nodes slightly
swollen; discoid cystoliths present, usually
denser at nodes. Probracts persistent, ovate,
rotund, oblong, rhomboid or broadly elliptic,
often cupular, base truncate, apex acute or
acuminate; margin entire, crenate or serrate
with 1-6 teeth, 2-14 mm long, 1.5-7 mm
wide, glands 2-8, glabrous or sparse minute
trichomes on both sides; discoid cystoliths
may be present. Tendrils unbranched or 2- or
3-branched. Leaves compound, digitate with
3-5 leaflets (juveniles simple or 3-foliolate,
cordate, ovate or elliptical, lobed or unlobed),
coriaceous; petioles 15-60 mm long, trichomes
sparse and minute, discoid cystoliths may be
present; central leaflets sessile or petiolules
up to 13 mm long; lamina elliptic, obovate,
elliptic-obovate or ovate-elliptic, 45-160 mm
long, 18-95 mm wide, base cuneate, rounded
or decurrent, apex acute or acuminate with a
soft mucro; margin serrate, remotely serrate
or undulate; 3-7 teeth, mucros or calluses
per side; lateral leaflets sessile or petiolules
up to 12 mm long, ovate or oblique-cordate,
50-170 mm long, 22-110 mm wide, lobed
or unlobed, base oblique, rounded, cuneate,
attenuate or cordate, apex acute or acuminate
with a soft mucro; margin serrate, remotely
serrate or undulate with 3-13 teeth, mucros
or callouses per side; upperside coarsely
scabrous, glabrous or with a few minute
trichomes along main veins especially near
base, numerous small translucent rosette¬

shaped cystoliths; underside scabrous or
smooth, glabrous or with a few translucent
trichomes near base, discoid cystoliths on
main veins; glands at base absent or up to
3, a few scattered usually present, diameter
0.6-1 mm. Male inflorescences with flowers
racemose or coaxillary with a solitary flower,
racemes 92-180 mm long with 3-14 flowers,
rachis thickened and 6-ribbed and with sparse
minute trichomes; peduncle 10-85 mm long,
3-5.25 mm wide, 6-ribbed; bracts obovate or
rhomboid, 25-40 mm long, 23-33 mm wide,
base truncate, apex obtuse or broadly obtuse,
margin fimbriate in upper half, venation
parallel, reticulation sparse, both sides
pubescent, glands scattered, gland diameter
0.5-1 mm. Male flowers: pedicels 3-5 mm
long; receptacle tube slender-funnelform,
10-angled or 10-ribbed, 38-46 mm long,
sparse minute trichomes; sepals subulate
or triangular, entire or 5-toothed, 22-38
mm long, 6-8 mm wide, green, 5-veined,
trichomes sparse and minute; corolla diameter
c. 50 mm (not including fimbrillae); petals
obdeltoid, 14-25 mm long, 15-20 mm wide,
white, pubescent. Female inflorescences
with flowers solitary or paired (one much
older). Female flowers: pedicel 10-19 mm
long; receptacle tube, salverform, 10-angled
or 10-ribbed, 40-47 mm long, glabrous
or glabrescent; sepals narrow-triangular,
entire, 5-veined, 11-15 mm long, green or
yellowish-green, clothed in minute trichomes;
corolla diameter 34-40 mm (not including
fimbrillae); petals obdeltoid, 14-18 mm long,
c. 9 mm wide, white, trichomes dense and
minute on both sides; stigma green; ovary
13-16 mm long, trichomes sparse and minute;
ovules several, in vertical rows, c. 1 mm long.
Fruit globose, ellipsoid, apex beaked, 10-
ribbed, 40-75 mm long, 50-65 mm wide,
red, glabrous or with a few trichomes at base;
mesocarp 11-14 mm wide, orange; pedicel 6-
36 mm long, 3-7 mm thick, trichomes sparse,
cystoliths absent or few to numerous; seeds
several, oblong or elliptical, flat or slightly
swollen, base rounded and may be flattened,
apex obtuse or acute, seeds 7-15 mm long,
5-8 mm wide, 2-4 mm thick, testa brown,
pulp dark green. Germination is hypogeal,
cotyledons straight. Figs. 1 & 6.

380

Austrobaileya 8(3): 364—386 (2011)

Fig. 6. . Trichosanthes pentaphylla. A. glandular probract at node. B. fruit, probract and tendril. C. adaxial view of
seeds. D. lateral view of seed. E. underside of leaf showing veins and glands. F. male inflorescence with glandular
bracts and bud. A-D from Cooper 2124 & Ford (CNS); E & F from Cooper 2134 & Jensen (CNS) (scales as indicated).
Del. W.T. Cooper.

Additional selected specimens (from 59 examined ):
Queensland. Cook District: 22 km NE of Bamaga, Feb
1994, Fell 4040 & Stanton (BRI); Lockerbie Scrub, Feb
1994, Cooper 814 (CNS); Bamaga, May 1981, Hyland
21136V (CNS); Merluna, Oct 1999, Hyland 16278
(CNS); Mowbray River Road, 3.5 km from highway, Jan
2000, Gray 7762 (CNS); Oak Beach, May 1974, Foley
431 (CNS); Kuranda, May 1993, Cooper 535 (CNS);
Kuranda Range Road, May 1976, Gray 163 (CNS); Bank
of Freshwater Creek, Freshwater, Nov 1936, Hunter s.n.
(CNS [NQNC2541); Aloomba, edge of Mulgrave River,
May 1998, Kitchener 73 (CNS); SFR310, Mar 1979, Gray
20088V (CNS); Royal Arch Tower, Chillagoe NP, Jun

2010, Cooper 2124 & Ford (CNS); Royal Arch Tower,
Chillagoe NP, Feb 2011, Cooper 2134 & Jensen (CNS);
Bank of North Johnstone River, Goondi near Innisfail,
Mar 1973, Colman s.n. (BRI [AQ9203]). North Kennedy:
Ingham, Jun 1997, Waterhouse 4431 (BRI); Townsville,
Jun 1936, Rowse s.n. (BRI [AQ310052]); Herbert River,
Euramo Station, Dansie s.n. (CNS [QRS119011]); Ayr,
Jan 1980, McGuire s.n. (BRI [AQ319650]); Home Hill
district, 1964, Wyatt s.n. (BRI [AQ310049]).

381

Cooper & de Boer, Trichosanthes in Australia

Distribution and habitat: Trichosanthes
pentaphylla is known from northeast
Queensland between Cape York and Home
Hill south of Townsville (Map 3). It mostly
occurs on riverbanks where it climbs into the
canopy and hangs along the forest edge in
wet lowland rainforest. It also occurs in the
Chillagoe-Mungana Caves N.P. at Chillagoe
where the habitat is deciduous vine thicket on
limestone.

Phenology: Flowers have been recorded in
January, February, March, April and May;
fruit has been recorded in January, February,
March, April, May, June and December.

Typification: A diverse range of specimens
were cited in the original description of this
species. These were widely distributed by
Mueller to herbaria in Europe with some
(but not all) having what are presumably
duplicates retained at MEL. We have selected
as lectotype of this name, a fertile collection
that is labelled as being from “Mt Grame”
and collected in June 1864, presumably by
Dallachy.

Other residual syntypes are as follows
- Australia: Queensland. North Kennedy
District: Rockingham Bay, s.dat., [probably
J.Dallachy s.n. for F.Mueller] (BR6605889,
photo!; K, photo!); Burdekin River, s.dat.,
[probably J.Dallachy s.n. for F.Mueller]
(MEL100121; K742661, photo!); Burdekin,
s.dat., [probably J.Dallachy s.n. forF. Mueller]
(K, photo!).

Notes: Trichosanthes pentaphylla specimens
from the Chillagoe area are rather variable
and could represent a new subspecies.
However, DNA analysis has not confirmed
this (de Boer & Cooper, unpublished). One
collection with male inflorescences is typical
of specimens from the lowland rainforests,
with a thickened rachis; bracts obovate or
rhomboid, margin fimbriate in upper half
and sparse reticulation. A second specimen
(Cooper 2101 & Ford ) from the same vicinity
has a slender rachis, bracts broadly ovate or
reniform, with an acute or acuminate apex,
margin denticulate not fimbriate, reticulate
venation and female receptacle tube deeply
ribbed. The morphology of this particular
specimen has not been included in the general
description.

Affinities: Telford (1982) suggested that
Trichosanthes pentaphylla may be conspecific
with T. trifolia (L.) Merr. (syn. T. wawrae).
However, Trichosanthes wawrae has narrowly
ovate probracts, 3-5 x 1.5-3 mm; suborbicular
juvenile leaves; adult leaves finely scabrous
above, margin entire or minutely sparsely
dentate, central leaflet base long-cuneate;
male raceme not thickened; bracts obovate or
oblong, 7-25 x 4-12 mm; sepals 4-6 mm long;
fruit pedicel 2-3 mm thick. T. pentaphylla has
probracts ovate, rotund, oblong, rhomboid
or broadly elliptic, 2-14 mm long, 1.5-7
mm wide; juvenile leaves cordate, ovate or
elliptical; adult leaves coarsely scabrous
above, margin serrate, remotely serrate or
undulate; central leaflet base cuneate, rounded
or decurrent; male raceme thickened; bracts
obovate or rhomboid, 25-40 mm long, 23-33
mm wide; sepals 22-38 mm long; fruit pedicel
3-7 mm thick.

Etymology: The specific epithet is derived
from the Greek penta- (five) and -phyllus (-
leaved) referring to the digitate compound
leaves, which often have 5 leaflets.

6. Trichosanthes morrisii W.E.Cooper
species nova a Trichosanthi pentaphyllae
similis, foliis simplicibus et sepalis margine
laciniato differt. Type: Australia: Northern
Territory. Kakadu National Park, near Gubara
Saddle, 28 December 2010, W.E.Cooper 2128,
I.Morris & R.Dempster (holo: CNS [2 sheets
+ spirit], iso: 8 sheets to be distributed to BR,
BRI, CANB, DNA, L, MO, P, UPS).

Trichosanthes sp. (D55403); Liddle et al.
(1994).

Trichosanthes sp. Kakadu (C.R. Dunlop
6639), Australian Plant Census (CHAH 2005);
Australian Plant Names Index, http://www.
anbg.gov.au/cgi-bin/apni Accessed February
2010 .

Dioecious trailing vine or liana to about c.
6 m, perennial, seasonally senescent. Stem
diameter to 20 mm, cream with numerous
black warts; young branchlets 7-ribbed with
cystoliths near nodes. Probracts persistent,
broadly elliptical, obovate or oblong; apex
acute or acuminate; margin denticulate,
crenate or serrate with 5-7 teeth, base

382

truncate or cuneate, 5-10 mm long, 2-4 mm
wide, adaxial surface with 4-7 glands, sparse
trichomes on both sides, cystoliths may be
present abaxially. Tendrils 2-4-branched.
Leaves simple, discolorous, membranous;
petioles 22-60 mm long, sparsely hairy,
discoid cystoliths present, sparse minute
trichomes may be present; lamina cordate,
3-5-lobed (juveniles cordate-deltoid, unlobed
or remotely 3-lobed), 80-170 mm long,
68-160 mm wide, base truncate, cuneate
or cordate, sinus deep and narrow or wide
(juveniles with shallow and wide sinuses),
apex acute or acuminate, usually with a soft
mucro; margin remotely denticulate with
17-24 mucros or teeth per side; upperside
scabrous, trichomes may be present on midrib
and main lateral veins; cystoliths numerous,
multicellular, rosette-shaped, diameter 0.3-1
mm; underside glabrous or a few trichomes
may be present near base; glands sparse near
base and towards apex, gland diameter 0.5-
1.25 mm; cystoliths discoid, sparse, scattered

Austrobaileya 8(3): 364-386 (2011)

along main veins. Male inflorescences with
flowers (from mature buds) racemose, usually
beside a solitary flower, racemes 140-240 mm
long, up to 11-flowered, peduncle 95-105 mm
long, rachis slender, trichomes sparse, discoid
cystoliths present; bracts persistent, rhomboid
or ovate, unlobed or shallowly lobed, 27-34
mm long, 17-24 mm wide, base attenuate,
cuneate or decurrent, apex acuminate; margin
denticulate, laciniate or fimbriate in upper
half only (entire in lower half); venation
palmate-pinnate, reticulation dense, both
sides with sparse minute trichomes denser
near margin towards apex, rarely with glands,
gland diameter c. 0.5 mm; aberrant leaf¬
like bracts may be present towards base of
inflorescence, ovate-cordate, 3-lobed, 26-57
mm long, 18-63 mm wide, base short cuneate
or decurrent, apex acute or acuminate, margin
denticulate, venation palmate-pinnate, 3-5-
veined (venation similar to leaves), reticulate,
trichomes as in main bracts. Male flowers:
pedicel on solitary flowers c. 40 mm long,

Fig. 7. Trichosanthes morrisii. A. male bract. B. seeds: left, lateral view; right, adaxial view. C. male sepal. D. leaf
underside with glands, tendril & male inflorescence. A, C, D from Cooper 2128, Morris & Dempster (CNS); B from
Dunlop 6626 (DNA) (scales as indicated). Del. W.T. Cooper.

383

Cooper & de Boer, Trichosanthes in Australia

sepals rhomboid, 3-veined, margin laciniate,
hairy, reticulate; petals not seen. Female
flowers: not seen. Fruit globose, apex beaked,

36- 55 mm long, 35-60 mm wide, orange to
red, probract persistent at base; mesocarp firm,
7-9 mm wide; pedicel up to 18 mm long and
5 mm wide, cystoliths present; seeds several,
flat, oblong, both ends obtuse, base slightly
bulbous, 11.2-12 mm long, 6.8-7.4 mm wide,

37- 4.2 mm deep, testa brown, suspended
in greenish-black pulp. Germination type
unknown. Fig. 7.

Additional selected specimens (from 8 examined ):
Australia: Northern Territory. Arnhem Land, SE Mt
Howship, Feb 1984, Dunlop 6626 (DNA); Arnhem Land,
SE Mt Howship, Feb 1984, Dunlop 6639 (DNA); Groote
Eylandt, Umbakumba, Mar 1988, Russell-Smith 5099 &
Lucas (DNA); Groote Eylandt, 8 km SW Umbakumba,
Jul 1987, Russell-Smith 2750 & Lucas (DNA); Groote
Eylandt, Jul 1987, Russell-Smith 2798 & Lucas (DNA);
Mt Brockman Outlier, Apr 1989, Russell-Smith 8056
(DNA); 1 km east of the Gubara Saddle, Kakadu N.P.,
Dec 2009, Cooper 2098 & Morris (CNS).

Distribution and habitat : Trichosanthes
morrisii is endemic to the Northern Territory
where it is currently known to occur in Kakadu
N.P., Arnhem Land and on Groote Eylandt
(Map 3). It inhabits Allosyncarpia-dominated
forest in gorges on sandstone escarpments
and in riparian evergreen vineforest on
coastal dunes. It appears to favour areas with
perennial water, which retain some moisture
through the drier seasons. Other species it co¬
occurs with are Remusatia vivipara (Roxb.)
Schott, Drynaria quercifolia (L.) J.Sm.,
Calophyllum sil Lauterb., Myristica insipida
R.Br., Tylophora benthamii Tsiang and
Desmos wardianus (F.M.Bailey) Jessup.

Phenology : Male inflorescences without
fully expanded flowers were collected in
December, and flowering probably occurs
between December and February; fruit has
been collected in February and April.

Affinities: Trichosanthes morrisii is closely
related to T. pentaphylla with similar juvenile
leaves and fruit. The former has simple
leaves, male bracts which are reticulate with
an acuminate apex, attenuate or cuneate base,
and lacinate sepals, whereas, T. pentaphylla
has compound leaves, male bracts are not
reticulate and have an obtuse apex, a truncate
base and entire sepals (excepting one aberrant

collection [Cooper 2101 & Ford ] in which the
bracts are ovate, reticulate and acute).

Etymology: Named for Northern Territory
naturalist, Ian Morris (1951—), in honour of
his extensive contribution to natural history
knowledge.

Acknowledgements

The authors thank the curators of the following
herbaria for allowing the examination of their
collections, either at the herbarium or through
loans or high resolution scans: A, BR, BRI,
CANB, CNS, DNA, E, K, KUN, L, MEL,
NSW, P, PERTH, S and UPS. Some specimen
data were sourced from Australia’s Virtual
Herbarium with permission of the Council of
Heads of Australasian Herbaria Inc.

The first author thanks Frank Zich &
Darren Crayn for support and access to CNS
herbarium; Andrew Ford, Rigel Jensen & Ian
Morris for much appreciated expertise and
companionship in the field; Bill Cooper for
the illustrations, Lyn Craven for the Latin
diagnosis, Steve Murphy for maps, Yumiko
Baba for translation; Sarah Legge, Keith
McDonald, Stephen McKenna, Hugh & Nan
Nicholson, Caroline Puente-Lelievre, Rupert
Russell and Jeremy Russell-Smith for various
contributions.

The first author is grateful for permits to
collect, which were issued by the Queensland
Department of Environment and Resource
Management as well as the Australian
Government (Kakadu National Park).

The second author was supported by
SIDA-SAREC grant SWE-2005-338.

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Map 1. Distribution of Trichosanthes cucumerina var. cucumerina (•) in Australia.

Map 2. Distribution of Trichosanthespilosa (A) and T. subvelutina (•) in Australia.

386

Austrobaileya 8(3): 364—386 (2011)

Map 3. Distribution of Trichosanthespentaphylla (A) and T. morrisii (•).

Five new species of Plectranthus L.Her. (Lamiaceae)
from New South Wales and Queensland

Paul I. Forster
Summary

Forster, P I. (2011). Five new species of Plectranthus L.Her. (Lamiaceae) from New South Wales
and Queensland. Austrobaileya 8(3): 387-404. Plectranthus caldericola P.I.Forst. from New South
Wales and P. bellus P.I.Forst., P. fragrantissimus P.I.Forst., P. insularis P.I.Forst. and P. venustus
P.I.Forst. from Queensland are described as new species. All species are illustrated together with
notes on their habitat, distribution, affinities and conservation status.

Key Words: Lamiaceae, Plectranthus, Plectranthus bellus, Plectranthus caldericola, Plectranthus
fragrantissimus, Plectranthus insularis, Plectranthus venustus, Australia flora. New South Wales
flora, Queensland flora, taxonomy, new species, conservation status

PI. Forster, Queensland Herbarium, Department of Environment & Resource Management, Brisbane
Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email:paul.forster@
derm.qld.gov.au

Introduction

The genus Plectranthus L.Her. is highly
diverse in Australia with at least 35 described
species (Blake 1971; Forster 1992,1994,1996,
1997b, 1999, 2008). As noted previously,
speciation in the genus has occurred where
populations occur in isolated areas such as
mountain peaks or ranges and where suitable
habitat exists on areas of exposed rock
pavements and outcrops. This suite of habitat
factors has resulted in the ‘islands on islands’
effect (Porembski et al. 2000) and is widely
recognized as being a driving force in genetic
diversity and speciation (Carlquist 1974;
Grant 1981; Hopper 2000; Seine et al. 2000;
Kruckeberg 2002). The swift maturation
of Plectranthus individuals from seed (one
growing season) with the potential for rapid
generation turnover enables speciation within
a given locality if a suitable niche is present and
once new genetic changes arise in a population.
These changes are perhaps most likely to
occur via genetic drift, rather than selection
for markedly different growth forms. This has
resulted in subtle differences in morphology
with a ‘non-adaptive radiation’ and a ‘high
lineage diversification rate’ (Savolainen &
Forest 2005), i.e. there are a lot of similar
appearing species in microhabitats that share
edaphic and topographic similarities and that
are often highly disjunct (Linder 2003).

Accumulation of a number of field
collections, together with a critical
examination of previously described taxa and
herbarium collections, now enable a further
five new species to be described, including
one long recognized, unnamed species in the
group (Forster 2002, 2007a; Bean & Forster
2007,2010). Two of these species are currently
known from single populations, whereas the
others are more widely distributed. All of
the newly described species occur in rocky
areas of pavements and slabs, on a range of
geological substrates.

Materials and methods

This work is based on herbarium specimen
collections in BRI, CANB (including CBG),
CNS (incorporating QRS) and MEL and
a large collection of live material that was
maintained in Brisbane in the 1990s. All
taxa were examined in habitat. Descriptive
terminology is similar to that previously used
(Forster 1994). Conservation assessments are
made using the IUCN (2001) criteria.

Accepted for publication 16 July 2011

388

Taxonomy

1. Plectranthus bellus RI.Forst., species nova

affinis P. spectabili S.T.Blake sed indumento
in caulibus et lamina foliorum e trichomatibus
glandulosis constanti (in illo absentibus) et
eglandulosis divaricatis (adversum antrorsis),
dentibus foliorum 10-19 in quoque latere (in
illo 7-12) et lobo superiore calycis oblongo-
ovato (non lanceolato-ovato) differens. Typus:
Queensland. Cook District: Daintree National
Park, Adeline Creek headwaters, 18 May 1999,
P.I.Forster PIF24610 & R.Booth (holo: BRI [2
sheets]; iso: MEL).

Erect to decumbent herb or subshrub to 100
cm high; foliage with very faint ‘lemon’
scent when crushed, clammy; non-glandular
and glandular trichomes uncoloured, sessile
glands 8-celled and yellow. Roots thickened
and tuberous to fibrous. Stems square, erect
to straggling, succulent, the lower parts up
to 12 mm diameter, pale green, upper parts
with persistent indumentum, non-glandular
trichomes dense and shaggy, divaricate,
6-16-celled up to 2 mm long, glandular
trichomes sparse, sessile glands absent.
Leaves discolorous, petiolate; petioles 4-18 x
2.5-3 mm, channelled on top, non-glandular
trichomes dense and shaggy, divaricate, 6-16-
celled up to 2 mm long, glandular trichomes
sparse, sessile glands absent; laminae ovate
to broadly ovate, ± succulent, 10-80 x 10-65
mm, crenate with 10-19 teeth up to 2 mm
long on each margin, widest above middle,
secondary teeth usually present; tip acute to
obtuse; base rounded to truncate; upper surface
silver-green, velutinous, veins impressed,
non-glandular trichomes dense, divaricate,
4-12-celled up to 1 mm long, glandular
trichomes absent or scattered to sparse,
sessile glands absent; lower surface silver-
white (due to the indumentum), velutinous,
veins raised, non-glandular trichomes dense
and shaggy, divaricate, 6-16-celled up to
2 mm long, glandular trichomes sparse,
sessile glands absent. Inflorescence up to 210
mm long, single or with 1-4 side branches,
pedunculate for 15-38 mm; axis square in
cross-section, non-glandular trichomes dense
and shaggy, divaricate, 4-16-celled up to 2
mm long, glandular trichomes sparse, sessile
glands absent; bracts broadly ovate, 3-4.2

Austrobaileya 8(3): 387-404 (2011)

x 2.8-3 mm, usually forming a tight coma,
non-glandular trichomes dense and shaggy,
divaricate, 6-10-celled up to 1 mm long,
glandular trichomes sparse, sessile glands
occasional; cymes sessile; verticillasters 5 or
6-flowered, up to 7 mm apart; pedicels 2-2.5
x c. 0.5 mm, non-glandular trichomes dense
and shaggy, antrorse to divaricate, 6-12-
celled up to 1mm long, glandular trichomes
sparse, sessile glands absent. Flower calyces
3-3.5 mm long, non-glandular trichomes
dense and shaggy, divaricate, 6-12-celled up
to 1 mm long, glandular trichomes sparse,
sessile glands absent. Corolla 8-10 mm
long, lilac to purple; tube 5-6 mm long,
curved at 90-100° 3-4 mm from the base,
slightly curved upwards, ± glabrous; upper
lobes suborbicular, reflexed, 2-2.2 x 1.2-1.5
mm, non-glandular trichomes occasional
to sparse, divaricate, 4-6-celled up to 0.8
mm long, glandular trichomes and sessile
glands absent; lateral lobes oblong, 1.8-2
x 1-1.2 mm, glabrous; lower lobe broadly
ovate, 4.5-5 x 3.8-4 mm, non-glandular
trichomes dense, divaricate, 4-6-celled up
to 1 mm long, glandular trichomes sparse,
sessile glands absent; filaments filiform, 9-10
x c. 0.1 mm, lilac, fused for c. 4 mm from
the base; anthers 0.6-0.7 x 0.4-0.5 mm; style
filiform, 10-11 x c. 0.2 mm, lilac, bifid for c.
0.3 mm. Fruit calyces 3-4 mm long; upper
lobe oblong-ovate, 1.8-2 x 0.8-1 mm; lateral
lobes lanceolate-falcate, 2.6-3 x 0.7-0.8 mm;
lower lobes lanceolate-falcate, 2.8-3 x 0.3-0.4
mm. Nutlets ± circular in outline, flattened to
somewhat convex, 0.7-0.8 mm long, 0.7-0.8
mm wide, 0.4-0.5 mm thick, glossy brown,
surface smooth. Fig. 1-3.

Additional specimens examined : Cook District:
Hann Tableland, track to Radar tower station. May
2006, Forster PIF31735 & McDonald (BRI); Hann
Tableland N.P., Jul 2004, McDonald KRM2982 & Bean
(BRI). Cultivated: Tolga (ex plant collected by G. &
N. Sankowsky from base of Hann Tableland), Jul 1995,
Forster PIF17166 (BRI); Mt Coot-tha Botanic Gardens
(ex plant [collector unknown] from Hann Tableland),
Mar 2007, Forster PIF32451 (BRI).

Distribution and habitat : Plectranthus
bellus is endemic to north-east Queensland
and has been recorded once from the western
edge of the Daintree N.R and at two localities
on the Hann Tableland. Sterile plants of
what appears to be this species were also

Forster, five new species of Plectranthus

389

Fig. 1. Plectranthus bellus. A. flowering stem x0.6. B. abaxial view of leaf xl. C. adaxial view of leaf xl. D. portion of
inflorescence with two verticillasters, buds and flowers x2. E. lateral view of flower x4. F. face view of flower x4. All
from Forster PIF17166 (BRI). Del. W. Smith.

390

observed at several additional localities in
the foothills of the Hann Tableland in May
2010. Plectranthus bellus is invariably found
on rock surfaces and crevices on small (30-
200 m 2 ) areas of granite pavements that form
irregular mosaics (‘islands’) within a greater
area of eucalypt woodlands (Fig. 2 & 3). Only
scattered perennial shrubs and small annual
grasses and forbs are found in association.

Notes : There are some morphological
similarities to the much taller growing
Plectranthus spectabilis (Forster 1996), that
has been found close by with at least two
populations within 10 km on upland parts of
the Hann Tableland. Observed morphological
differences between the two species are

Austrobaileya 8(3): 387-404 (2011)

summarized in Table 1; however, the most
important differences relate to the overall
colour of the plants (due to a combination
of epidermis colour and indumentum cover)
(Fig. 3), together with the form of the flowers.
Other species of Plectranthus that also occur
on the Hann Tableland, but were not observed
to be locally sympatric with P. bellus , include
P. apreptus S.T.Blake and P. mirus S.T.Blake.

Plectranthus bellus is a quite spectacular
appearing plant and has considerable potential
in amenity horticulture and landscaping on
account of its attractive foliage, upright form
and the ability to grow in a range of soil types
in cultivation. Clones are self sterile, further
adding to its attraction as an amenity plant.

Fig. 2. Plectranthus bellus. Habitat at Hann Tableland (population voucher: Forster PIF31735 & McDonald [BRI])
Grasses in background are predominantly exotic species. Photo: P.I. Forster

Forster, five new species of Plectranthus 391

Table 1. A comparison of selected morphological character states for Plectranthus bellus
and P. spectabilis

Character state

P. bellus

P. spectabilis

life form

perennial, herb or subshrub to 1 m tall

annual or biennial, herb or
subshrub to 1.6 m tall

indumentum on
stems and leaf
laminae

non-glandular trichomes dense
and shaggy, divaricate; glandular
trichomes generally present

non-glandular trichomes
sparse to dense, antrorse;
glandular trichomes absent

number of teeth
on leaf laminae
margins

10-19

7-12

inflorescence

up to 210 mm long with 1-4 branches
from near the base; non-glandular
trichomes divaricate

up to 320 mm long with 1-11
branches from near the base
or further up the axis; non
glandular trichomes antrorse

calyx indumentum

divaricate

antrorse

upper lobe of calyx

oblong-ovate, 1.8-2 x 0.8-1 mm

lanceolate-ovate, 2.3-4 x
0.8-1.3

Fig. 3. Plectranthus bellus. Habit of individual plant (population voucher: Forster PIF31735 & McDonald [BRI]). Photo:
P I. Forster

392

Conservation status: This species is currently
known from at least three populations with a
known area of occupancy of less than 5 km 2 .
The population in the Daintree N.P. occurs in
an area that is more or less pristine with little
evidence of weed infestation. The populations
at the Hann Tableland are besieged by invasive
alien weeds that have been spread widely
over the tableland by grazing cattle and
uncontrolled wildfires prior to gazettal of the
area as National Park. These small islands of
granite pavement are under direct and current
threat from weeds such as Andropogon
gay anus Kunth, Bidens pilosa L., Lantana
camara L., Melinis minutiflora PBeauv., M.
repens (Willd.) Zizka and Praxelis clematidea
R.M.King & H.Rob. that compete for the
scarce resources in these habitats and greatly
increase fuel loads both on the pavements and
in adjacent vegetation. These Plectranthus
species are unable to survive hot fires with the
plants being killed outright or being forced to
resprout from subterranean rootstocks. Due
to the shallow soils on these pavements, the
heat and scorch from any fires often tends to
also kill the rootstocks of Plectranthus. The
infrequent occurrence of this species on small
pavements (<20 m 2 ) (Fig. 2) may be regulated
by fires that occur not necessarily only on the
pavement, but in adjacent grassy vegetation.

Map 1 . Distribution of Plectranthus bellus in north¬
east Queensland. Shaded area on map indicates
conservation reserves (National Parks, Forest Reserves
and Conservation Parks).

Austrobaileya 8(3): 387-404 (2011)

The recommended conservation status
for Plectranthus bellus is Vulnerable (criteria
B2,a,b(i,ii,iii,iv,v), D2) based on the IUCN
(2001) categories and criteria.

Etymology: The specific epithet is formed
from the Latin word bellus (beautiful) and
alludes to the pleasing form of this plant.

2. Plectranthus caldericola P.I.Forst., species
nova a P. graveolenti R.Br. foliis caulibusque
siccis non viscidis leviter citriodoris (in illo
humidis viscidis et praeditis odore anti-
septico gravi), foliis dentibus paucioribus
praeditis (14-18 non 20-42) et apicibus
rotundatis (in illo acutis usque ad obtusis),
verticillastris 2-14-floris (adversum 12-15) et
floribus pedicellis brevioribus praeditis (0.8-
1.8 mm non 2-4.5 mm) differt. Typus: New
South Wales. The Pinnacle, 13 km W of Mt
Warning, 6 April 1994, P.I. Forster PIF15093
& G.Leiper (holo: BRI [3 sheets ± spirit]; iso:
MEL, NSW).

Subshrub to 50 cm high; foliage with faint
Temon’ scent when crushed, not clammy;
non-glandular and glandular trichomes
uncoloured to silver, sessile glands 8-celled
and orange. Roots thickened and tuberous.
Stems square, erect to straggling, fleshy,
the lower parts up to 15 mm diameter, dark
purple, non-glandular trichomes dense and
shaggy, divaricate, 6-10-celled up to 3 mm
long, glandular trichomes sparse, sessile
glands scattered. Leaves discolorous, petiolate;
petioles 5-33 x 2-2.5 mm, channelled on top,
non-glandular trichomes sparse, divaricate,
6-10-celled up to 3 mm long, glandular
trichomes sparse, sessile glands scattered;
laminae broadly ovate to almost lobate, fleshy,
10-50 x 10-50 mm, crenate with 7-9 teeth up
to 3 mm long on each margin, widest above
middle, secondary teeth absent; tip rounded;
base truncate, cordate or lobate; upper surface
dark-green, veins impressed, non-glandular
trichomes sparse, divaricate, 2-8-celled up
to 1 mm long, glandular trichomes scattered,
sessile glands sparse; lower surface pale-green,
veins raised, non-glandular trichomes dense,
divaricate, 2-10-celled up to 2 mm long,
glandular trichomes sparse, sessile glands
dense. Inflorescence up to 320 mm long,
comprising 1-5 pedunculate branches from
near the base; axis square in cross-section,

Forster, five new species of Plectranthus

393

Fig. 4. Plectranthus caldericola. A. flowering stem x0.4. B. abaxial view of leaf xl. C. adaxial view of leaf xl. D. portion
of inflorescence with two verticillasters, buds and flower x3. E. lateral view of flower x6. F. face view of flower x6. G.
lateral view of fruiting calyx x6. All from Forster PIF15093 & Leiper (BRI). Del. W. Smith.

394

non-glandular trichomes sparse, divaricate,
6-10-celled up to 5 mm long, glandular
trichomes dense, sessile glands scattered;
bracts broadly-ovate to obovate, 3-5 x 2.5-4
mm, not forming a coma but approaching one,
non-glandular trichomes dense, divaricate, 6-
10-celled up to 3 mm long, glandular trichomes
absent, sessile glands scattered; cymes sessile;
verticillasters 2-14-flowered, 5-15 mm
apart; pedicels 0.8-1.8 x c. 0.2 mm diameter,
non-glandular trichomes dense, divaricate,
6-10-celled up to 2 mm long, glandular
trichomes absent, sessile glands scattered.
Flower calyces 1.8-2 mm long, non-glandular
trichomes dense and shaggy, divaricate,
6-10-celled up to 3 mm long, glandular
trichomes absent, sessile glands dense. Corolla
6-9 mm long, dark-purple; tube 3-4 mm long,
± straight or weakly curved at 160-170° 2-2.5
mm from base, slightly inflated upwards,
glabrous; upper lobes orbicular, erect, 1-1.8
x 1-2 mm, non-glandular trichomes dense,
divaricate, 2-4-celled up to 0.6 mm long,
glandular trichomes absent, sessile glands
scattered; lateral lobes oblong, 1-1.5 x 0.5-0.6
mm, glabrous; lower lobe ± orbicular, 3-4.5 x

3.5- 4.5 mm, non-glandular trichomes sparse,
divaricate, 2-4-celled up to 0.3 mm long,
glandular trichomes absent, sessile glands
scattered; filaments filiform, 7-11 x c. 0.2 mm,
lilac, fused for 3-4 mm from base; anthers c.
0.5 x 0.3 mm; style filiform, 8-9 mm long,
lilac, bifid for c. 0.8 mm. Fruit calyces 3-3.5
mm long; upper lobe orbicular-ovate, 1.2-1.6 x

1.5- 1.7 mm; lateral lobes lanceolate, 1.2-1.5 x

O. 9-1 mm; lower lobes lanceolate-falcate, 1.2-
1.5 x 0.8-1 mm. Nutlets ± circular in outline, ±
flattened, 0.6-0.7 mm long, 0.6-0.7 mm wide
and c. 0.5 mm thick, glossy brown, surface
smooth. Fig. 4.

Additional specimens examined: New South Wales.
The Pinnacle, Jan 1993, McDonald s.n. (BRI, NSW).

Distribution and habitat : Plectranthus
caldericola is known only from The Pinnacle
in the Border Ranges of northern New South
Wales. Plants grow on rock outcrops and
pavements formed of porphyritic basalts
between 920 and 1000 m with a shrubland
dominated by Leptospermum polygalifolium
Salisb. Other species of Plectranthus that
occur sympatrically are P. graveolens and

P. parviflorus Willd.

Austrobaileya 8(3): 387-404 (2011)

Notes : This species was first brought to my
attention by Graham McDonald in April
1994. Plectranthus caldericola is distinctive
in the very broad leaves that are sometimes
almost lobate, the shaggy indumentum on the
foliage and the very hairy calyces and very
short pedicels. It differs from Plectranthus
graveolens in the non-clammy foliage with
a faint Temon’ scent (versus clammy with a
strong antiseptic or foetid scent), leaves with
fewer teeth to the lamina (14-18 versus 20-42)
and rounded tips (versus acute to obtuse), the
2-14 flowered verticillasters (versus 12-15)
and the flowers with short pedicels (0.8-1.8
mm versus 2-4.5 mm).

The possibility that this species is an
example of an allotetraploid derived from an
initial hybridization between Plectranthus
graveolens and P. parviflorus needs
investigation. Plectranthus caldericola is not
typical of what one would expect from an FI
hybrid in terms of its morphology and ‘wild’
hybrids of Plectranthus have proven to be
rarely encountered (Blake 1971: 29; Forster
1991). Further localities should be sought for
this species in adjacent areas and the status of
somewhat similar plants (perhaps also from
putative hybridization events) at the North
Obelisk in New South Wales and Buchanan’s
Fort in Queensland requires further work.

Conservation status : Plectranthus caldericola
is presently known only from the type locality
where it is a very common plant. Although
this locality is within National Park, the
plants may be subjected to trampling from
bushwalkers and from the occasional wildfire.
The recommended conservation coding is
Vulnerable based on the criterion D2 (IUCN
2001 ).

Etymology : The specific epithet is derived
from calderus (a caldera) and cola (loving)
and refers to the occurrence of this species on
part of the Mt Warning caldera.

3. Plectranthus fragrantissimus P.I.Forst.,
species nova affinis P. suaveolenti S.T.Blake,
sed lamina foliorum minore angustiore
(lanceolato-ovata usque ovata, 10-40 x
6-25 mm non ovata usque late ovata, 30-
75 x 20-60 mm), florum pedicellis multo
longioribus (4-5.5. mm adversum 1.5-4 mm),

Forster, five new species of Plectranthus

395

Map 2. Distribution of Plectranthus caldericola ★,
P. fragrantissimus • and P. insularis A in southern
Queensland and northern New South Wales. Shaded area
on map indicates conservation reserves (National Parks,
Forest Reserves and Conservation Parks).

calyce lobis inferioribus lobos laterales plus
minusve aequantibus et lobis superioribus
ovatis (non subcircularibus) differens. Typus:
Queensland. Port Curtis District: State
Forest 316, Kroombit Tops, 16 February 1995,
P.I.Forster PIF16246 (holo: BRI [2 sheets +
spirit]; iso: MEL, NE).

Low growing herb to 30 cm high; foliage
with sweetly aromatic scent when crushed,
clammy; non-glandular and glandular
trichomes uncoloured, sessile glands absent.
Roots thickened, fibrous. Stems square, erect
to straggling, fleshy, the lower parts up to 4
mm diameter, pink-purple, non-glandular
trichomes sparse to dense, retrorse, 2-6-celled
up to 0.3 mm long, glandular trichomes absent
or scattered to sparse. Leaves discolorous;
petioles 2-19 x 1-1.7 mm, channelled on
top, non-glandular trichomes dense, retrorse,
2-6-celled up to 0.8 mm long, glandular
trichomes sparse; laminae lanceolate-ovate to
ovate, fleshy to succulent, 10-40 x 6-25 mm,
crenate with 7-14 teeth up to 2 mm long on
each margin, widest above middle, secondary
teeth sometimes present; tip acute; base

cuneate to truncate; upper surface dark-green
to silver-green, veins impressed, non-glandular
trichomes sparseto dense, antrorseto divaricate,
6-8-celled up to 1 mm long, glandular
trichomes sparse to dense; lower surface silver-
green, veins raised, non-glandular trichomes
dense (particularly on veins), retrorse, 6-
8-celled up to 1 mm long, glandular trichomes
sparse to dense. Inflorescence up to 230 mm
long, usually unbranched (rarely with two
side branches); axis square in cross-section,
non-glandular trichomes sparse to dense,
retrorse, 2-6-celled up to 0.2 mm long,
glandular trichomes scattered to sparse; bracts
obovate, 1.8-2 x 1.4-2 mm, not forming a
coma, non-glandular trichomes sparse to
dense, antrorse to divaricate, 2-8-celled up
to 0.6 mm long, glandular trichomes sparse;
cymes sessile; verticillasters 6-14-flowered,
4-20 mm apart; pedicels 4-5.5 x c. 0.2 mm
diameter, non-glandular trichomes sparse to
dense, antrorse to divaricate, 2-4-celled up
to 0.2 mm long, glandular trichomes scattered
to sparse. Flower calyces 2.5-3.5 mm long,
non-glandular trichomes scattered to sparse,
antrorse to divaricate, 2-6-celled up to 0.5
mm long, glandular trichomes sparse to dense.
Corolla 9-10 mm long, lilac; tube 5-5.5 mm
long, weakly curved at 10-20° c. 2 mm from
base, slightly inflated upwards, non-glandular
trichomes scattered, divaricate to retrorse, 2-4-
celled up to 0.2 mm long, glandular trichomes
absent; upper lobes suborbicular, erect, 1.8-2 x
1.8-2 mm, non-glandular trichomes scattered
to sparse, divaricate, 2-4-celled up to 0.2
mm long, glandular trichomes scattered to
sparse; lateral lobes oblong, 1.3-1.8 x 0.6-0.8
mm, glabrous or as in upper lobes; lower lobe
oblong-ovate, 4.8-5 x 3.8-4 mm, indumentum
as in upper lobes; filaments filiform, 6-8 x c.
0.2 mm, lilac, fused for 3-4 mm from base;
anthers c. 0.4 x 0.3 mm; style filiform, 8-9 mm
long, lilac, bifid for c. 0.3 mm. Fruit calyces

3.5- 4.8 mm long; upper lobe ovate, 1.5-2 x
1.7-2 mm; lateral lobes lanceolate, 1.5-2 x
0.8-0.9 mm; lower lobes lanceolate-falcate,

1.5- 2.2 x c. 0.5 mm. Nutlets ± circular in
outline, ± flattened, 0.7-0.9 mm long, 0.7-0.9
mm wide, 0.4-0.5 mm thick, glossy dark
brown, surface smooth. Fig. 5.

Additional specimens examined : Queensland. Port
Curtis District: S.F. 316 Kroombit Tops, Feb 1995,

396

Austrobaileya 8(3): 387—404 (2011)

Fig. 5. Plectrantlius fragrantissimus. A. flowering stem *1. B. abaxial view of leaf x2. C. adaxial view of leaf x2. D.
portion of inflorescence with two verticillasters, buds and flowers x4. E. lateral view of flower x6. F. face view of flower
x9. G. pedicel and calyx x9. All from Forster PIF29770 & Halford { BRI). Del. W.Smith.

Forster, five new species of Plectranthus

Forster PIF16252 (BRI, MEL, NSW); loc. cit., Feb 1995,
Forster PIF16254 (BRI); Kroombit Tops F.R., near radio
tower, Nov 2003, Forster PIF29770 & Halford (BRI,
MEL); Kroombit Tops, S.F. 316, Mar 1995, Thompson
BIL156 et al. (BRI); Kroombit Creek, Kroombit Tops
N.P., Feb 2009, Mathieson MTM307 (BRI); top of
range. Tableland Road, Kroombit Tops N.P., Dec 2010,
Mathieson MTM980 (BRI).

Distribution and habitat: Plectranthus
fragrantissimus is endemic to Kroombit Tops
National Park (formerly State Forest or Forest
Reserve) where it occurs in a series of locally
disjunct populations that may represent
subpopulations of one large metapopulation.
The plants occur at altitudes between 800
and 900 m on large outcrops or pavements
of metamorphosed quartzose sandstone that
are scattered within woodland dominated by
Eucalyptus eugenioides Sieber ex Spreng.,
E. longirostrata (Blakely) L.A.S.Johnson &
K.D.Hill and E. montivaga A.R.Bean. Typical
Plectranthusparviflorus is also widespread at
Kroombit Tops.

Notes: This species has obvious affinities
to Plectranthus suaveolens, from which it
is disjunct by about 450 km with the most
northerly recorded population of that species
west of Mt Huntley in the Main Range
near the Queensland/New South Wales
border. Plectranthus suaveolens occurs
on granophyre (Mt Barney), trachyte (Mt
Ballow), rhyolite (Mt Ernest) or granite
(Granite Belt and northern New South Wales),
but has not yet been recorded from sandstone.
As with Plectranthus suaveolens there is an
absence of sessile glands on the foliage of
P. fragrantissimus\ however, the two species
differ by the latter with smaller, narrower leaf
laminae (lanceolate-ovate to ovate, 10-40
x 6-25 mm versus ovate to broadly ovate,
30-75 x 20-60 mm), much longer flower
pedicels (4-5.5 mm versus 1.5-4 mm), the
calyx with lower lobes much the same length
as the lateral lobes (versus much longer) and
the ovate upper lobes (versus subcircular).

Plectranthus fragrantissimus is the
third angiosperm taxon endemic to the
Kroombit Tops tableland, the others being
Bulbophyllum weinthalii subsp. striatum
D.L. Jones and Parsonsia kroombitensis
J.B Williams. The tableland is also botanically
notable for a number of significant species

397

disjunctions (northern limits, indicated *) or
for geographically restricted species with a
significant proportion of their populations
being present there, including *Amperea
xiphoclada (Sieber ex Spreng.) Druce var.
xiphoclada, Astrotricha brachyandra
A.R.Bean, Boronia palasepala Duretto,
*Bossiaea rupicola A.Cunn. ex Benth.,
*Bulbine vagans E.M.Watson, Bulbophyllum
globuliforme Nicholls, *Callicomaserratifolia
Andrews, *Ceratopetalum apetalum
D.Don, Chiloglottis sylvestris D.L. Jones
& M.A.Clem., *Lasiopetalum ferrugineum
Sm. var. ferrugineum, *Maytenus silvestris
Lander & L.A.S.Johnson, Myrsine ireneae
subsp. curvata Jackes, *Persoonia volcanica
P.H.Weston & L.A.S.Johnson, *Thismia
rodwayi F.Muell., Triplarina volcanica subsp.
borealis A.R.Bean and Zieria distans Duretto
& RI.Forst. Consequently Kroombit Tops is
often regarded as an important Pleistocene
refugial area for both plant communities and
species (e.g. Rozefelds & Barnes 2001) and
can be considered a cooler upland ‘island’
analogous with the ‘sky island’ concept
applied elsewhere (Heald 1951; Watling &
Donnelly 2006). The small number of endemic
plants does not justify it being considered
as even a minor centre of endemism ( sensu
Crisp et al. 2001); however, when combined
with an increasing number of vertebrate and
invertebrate animals that are being recognized
as endemic to the area (e.g. Ponniah &
Hughes 2004), it easily qualifies as such. A
logical hypothesis is to regard Plectranthus
fragrantissimus and P. suaveolens as sister
taxa originating from an allopatric speciation
event following ‘stranding’ of the lineage on
‘sky islands’ at Kroombit Tops and the New
England Tableland and Border Ranges/Scenic
Rim of southern Queensland and adjacent
New South Wales.

Conservation status: Plectranthus

fragrantissimus is abundant at most of its
recorded localities; however, the overall area
of occurrence and occupancy is very small.
The recommended conservation status is
Vulnerable based on the criterion D2 (IUCN
2001 ).

Etymology: The specific epithet is derived
from the Latin fragrans (fragrant) and -issima

398

(most) and alludes to the sweetly smelling
foliage of this plant.

4. Plectranthus insularis RI.Forst. species
nova a P. intraterranea S.T.Blake lamina
foliorum lanceolata, lanceolato-ovata vel
oblanceolata (in illo ovata usque late ovata),
dentibus foliorum longitudine plus quam 1/3
partes latitudinis laminae ad medianum ejus,
et calyce fructificanti majore (5.8-6.5 mm
non 3.8-5.5 mm) differt. Typus: Queensland.
Darling Downs District: Mingimarny State
Forest 131, Pine Hill, 20 km S of Millmerran,
28 April 1995, P.I.ForsterPIF16468 & S.J.Figg
(holo: BRI [2 sheets + spirit]; iso: MEL).

Low multistemmed herb to 50 cm high; foliage
with faint aromatic scent, not clammy; non-
glandular and glandular trichomes clear, sessile
glands 8-celled and red. Roots thickened-
tuberous, but not ‘carrot’-like. Stems square,
erect to decumbent, fleshy with the lower
parts slightly woody and up to 5 mm diameter,
green to pink-purple, non-glandular trichomes
dense, retrorse, 2-4-celled up to 0.1 mm long,
glandular trichomes absent, sessile glands
dense. Leaves discolorous, petiolate; petioles
3-8 x 1-1.8 mm, grooved on top, non-glandular
trichomes dense, retrorse, 2-4-celled up to
0.2 mm long, glandular trichomes scattered,
sessile glands dense; laminae lanceolate,
lanceolate-ovate or oblanceolate, fleshy, 15-60
x 5-28 mm, markedly crenate with 5-8 teeth
1.5-5 mm long on each margin, 2-6 mm wide
at base, secondary teeth absent; tip acute;
base attenuate; upper surface pale green,
veins impressed, non-glandular trichomes
sparse, antrorse, 4-8-celled up to 2 mm long,
glandular trichomes sparse, sessile glands
sparse; lower surface pale green, veins raised,
non-glandular trichomes sparse, retrorse, 2-8-
celled up to 1 mm long (usually much shorter),
glandular trichomes scattered, sessile glands
dense. Inflorescence up to 180 mm long,
single-stemmed; axis square in cross-section,
non-glandular trichomes dense, retrorse, 2-4-
celled up to 0.1 mm long, glandular trichomes
scattered, sessile glands sparse; cymes sessile;
verticillasters 10-12 flowered, up to 22 mm
apart; bracts lanceolate-ovate, 2.5-3 x c.
1.5 mm, not forming a coma, non-glandular
trichomes sparse, antrorse, 2-4-celled to 0.2
mm long, glandular trichomes absent, sessile

Austrobaileya 8(3): 387-404 (2011)

glands sparse; pedicels 4.5-5 x c. 0.3 mm,
non-glandular trichomes dense, divaricate,
2-celled up to 0.1 mm long, glandular
trichomes dense, sessile glands scattered.
Flower calyces 3.5-4 mm long, non-glandular
trichomes sparse, divaricate, 2-4-celled up
to 0.1 mm long, glandular trichomes absent,
sessile glands dense. Corolla 10-11 mm long,
purple; tube 2-2.5 mm long, weakly curved at
30-45° 2-2.5 mm from base, barely inflated
upwards, glabrous; upper lobes subcircular,
reflexed, 2.2-2.5 x 2.2-2.5 mm, non-glandular
trichomes sparse, divaricate to retrorse, 2-4-
celled up to 0.1 mm long, glandular trichomes
absent, sessile glands dense; lateral lobes
oblong, 2-2.2 x 0.9-1 mm, glabrous; lower
lobe orbicular-ovate, 5-5.5 x 4.5-5.5 mm, non-
glandular trichomes scattered, divaricate, 2-
celled up to 0.1 mm long, glandular trichomes
absent, sessile glands sparse; filaments
filiform, 12-13 x c. 0.3 mm, lilac, fused for c.
4 mm from base; anthers 0.4-0.5 x c. 0.3 mm;
style filiform, 13-14 x 0.2-0.3 mm, lilac, bifid
for c. 0.5 mm. Fruit calyces 5.8-6.5 mm long;
upper lobe broadly ovate, 2.3-3.1 x 2.2-2.8
mm; lateral lobes lanceolate-falcate, 2.5-3 x
0.5-0.7 mm; lower lobes lanceolate-falcate, 3-
3.5 x 0.3-0.5 mm. Nutlets ± circular in outline,
convex, 0.8-0.9 mm long, 0.8-0.9 mm wide, c.

O. 6 mm thick, glossy brown, surface smooth.
Fig. 6.

Additional specimens examined: Queensland. Darling
Downs District: Pine Hill S.F., c. 22 km S ofMillmerran,
Nov 1997, Grimshaw PG2925 & van Baalen (BRI); Pine
Hill, Mingimarny S.F. 131,20 km S ofMillmerran, May
2000, Menkins ILM96 (BRI).

Distribution and habitat : Plectranthus
insularis is currently known only from the
type locality in an isolated State Forest south of
Millmerran in south-east Queensland where it
is the only species of this genus present. This
locality is unusual in being an outlier of granite
(with pavements and outcrops) supporting low
eucalypt woodland or shrubland surrounded
by dissimilar geology (mainly heavy clay
soils) that used to support brigalow open
forest prior to conversion to agricultural land.
The granite appears similar to that found
near Texas and Stanthorpe; however, the
only species of Plectranthus found in those
areas are P. graveolens, P. parviflorus and

P. suaveolens.

Forster, five new species of Plectranthus

Notes: Plectranthus insularis is perhaps one
of the most distinctive species in the genus
in Australia on account of the extreme lobing
(in comparison with the overall leaf laminae
size) with the marginal teeth commonly up to
5 mm long. Suggested affinities are somewhat
difficult to deduce based purely on morphology,
but include Plectranthusparviflorus (which is
closest geographically) and P. intraterraneus

399

(which is markedly disjunct with the nearest
locality in western Queensland, but perhaps
closest on morphological similarity). A
comparison of morphological character states
for these three species is given in Table 2 with
notable distinctions for Plectranthus insularis
being the leaf laminae shape, floral bract
shape and the length of the fruiting calyx.

Table 2. Comparison of some morphological character states for Plectranthus insularis ,
P. intraterraneus and P. parviflorus

Character state

P. insularis

P. intraterraneus

P. parviflorus

Habit

Erect to decumbent herb
to 50 cm high

Erect subshrub to 1
m high

Erect to decumbent
herb to 70 cm high

Root system

thickened tuberous

‘carrot’ tuberous

distinct tuber at
stem base with
fibrous roots

Foliage

faintly scented, not
clammy

sweetly scented,
clammy

faintly scented, not
clammy

Lamina shape

lanceolate, lanceolate-
ovate or oblanceolate

ovate to broadly ovate

ovate to oblong-
ovate; more rarely
± orbicular or
truncate-ovate

Leaf teeth

markedly crenate;

5-8 pairs with the teeth
length being over 1/3 of
the lamina width at its
midpoint

crenate to dentate -
crenate; 4-9 pairs
with the teeth length
being less than 1/3 of
the lamina width at its
midpoint

crenate to dentate-
crenate; 4-8
(-12) with the teeth
length being less
than 1/3 of the
lamina width at its
midpoint

Marginal teeth
length (mm)

1.5-5

1-4

0.8-2 (-4)

Number of
flowers per
verticillaster

10-12

6-10

6-10

Floral bracts

lanceolate-ovate, 2.5-3
mm long

broadly ovate to
circular ovate, 1.2-2.7
mm long

broadly ovate to
obovate, 1.1-2.5
mm long

Fruiting calyx
length (mm)

5.8-6.5

3.8-5.5

37-5.5

Conservation status: State Forest 131 is an
isolated pocket of intact vegetation within a
much greater area of cleared land. The granite
geology of this State Forest is also unique
within the context of the local Millmerran
area. Plectranthus insularis is locally very

common on pavement areas within the State
Forest, particularly at the western end. The
plants have been observed to either die, or
shrivel back to the rootstocks during periods
of drought.

400

Austrobaileya 8 ( 3 ): 387—404 ( 2011 )

Fig. 6. Plectranthus insularis. A. flowering stem x0.4. B. abaxial view of leaf xl.5. C. adaxial view of leaf xl.5. D.
portion of inflorescence with two verticillasters, buds and flowers x2. E. lateral view of flower x4. F. face view of flower
x4. All from Forster PIF16468 & Figg (BRI). Del. W. Smith.

The recommended conservation status for
Plectranthus insularis is Vulnerable (criterion
D2) based on the IUCN (2001) categories
and criteria. This species is known from a
single population (or single metapopulation
with subpopulations) with a very restricted
area of occupancy (<20 km 2 ) that is prone to
stochastic events in the short to long term.

Etymology : The specific epithet is derived
from the Latin adjective insularis (pertaining
to islands) and alludes to the occurrence of
this species on an island of granite pavement,
further surrounded by a ‘sea’ of cleared land.

Forster, five new species of Plectranthus

5. Plectranthus venustus P.I.Forst., species
nova affinis P. gratae S.T. Blake sed
caulibus praeditis trichomatibus antrorsis
non-glandularibus (adversum divaricatis),
foliis numero dentium marginalium majore
(22-33 in latere quoque non 13-15), bracteis
inflorescentiae obovatis quae comam format
(adversum ovatis et comam non formans) et
floris pedicellis multo brevioribus (0.5-1 mm
non 2.5-3 mm) differens. Typus: Queensland.
Cook District: Kennedy Hill Gorge, 21
June 1989, P.I. Forster PIF5388 & M.C. Tucker
(holo: BRI [1 sheet + spirit]; iso: CNS, MEL).

Plectranthus sp. (Restoration Island J. Le
Cussan 285); Forster (1997: 98); Forster
(2002: 96); Bean & Forster (2007: 95); Bean
& Forster (2010: 90).

Prostrate to semi-erect herb or subshrub to 40
cm high; foliage with very faint ‘lemon’ scent
when crushed, not clammy; non-glandular and
glandular trichomes uncoloured to purplish,
sessile glands 8-celled and orange-red. Roots
fibrous. Stems square, erect to straggling,
succulent, the lower parts up to 10 mm
diameter, green to pink, upper parts with
indumentum, non-glandular trichomes sparse,
retrorse, 6-10-celled up to 1.2 mm long,
glandular trichomes absent, sessile glands
absent. Leaves discolorous, petiolate; petioles
2-16 x 2.5-4 mm, slightly channelled on top,
non-glandular trichomes dense, divaricate,
6-12-celled up to 1.5 mm long, glandular
trichomes absent, sessile glands scattered to
dense; laminae lanceolate to narrow-ovate, ±
succulent, 26-90 x 14-50 mm, crenate with
22-33 teeth up to 1 mm long on each margin,
widest above middle, secondary teeth usually
present; tip acute; base cuneate to rounded;
upper surface silver-green, veins impressed,
velutinous, non-glandular trichomes sparse,
antrorse to divaricate, 6-10-celled up to 0.8
mm long, glandular trichomes absent, sessile
glands absent; lower surface silver, veins
raised, velutinous, non-glandular trichomes
dense, divaricate, 6-12-celled up to 1.5 mm
long, glandular trichomes absent, sessile glands
scattered to dense. Inflorescence up to 150 mm
long, with 3-5 pedunculate branches from
near the base; axis square in cross-section,
non-glandular trichomes scattered, retrorse,
6-10-celled up to 2 mm long, glandular

401

trichomes absent, sessile glands absent; bracts
obovate to broadly-ovate, 2-3 x 2.4-3.2
mm, usually forming a coma, non-glandular
trichomes dense, antrorse, 6-8-celled up to 1.8
mm long, glandular trichomes absent, sessile
glands scattered; cymes sessile; verticillasters
18-22-flowered, up to 10 mm apart; pedicels
0.5-1 x 0.2-0.3 mm, non-glandular trichomes
dense, divaricate, 6-10-celled up to 1.8 mm
long, glandular trichomes absent, sessile
glands scattered. Flower calyces 2.2-2.8
mm long, non-glandular trichomes dense,
divaricate, 4-10-celled up to 2 mm long,
glandular trichomes absent, sessile glands
sparse. Corolla 10-12.5 mm long, pale-lilac;
tube 4.8-5.5 mm long, curved at 100-120°
1.7-2 mm from base, slightly inflated upwards,
± glabrous; upper lobes suborbicular, reflexed,
2-2.2 x 1.8-2.2 mm, non-glandular trichomes
sparse, divaricate, 4-6-celled up to 0.7 mm
long, glandular trichomes absent, sessile
glands sparse; lateral lobes oblong, 1.5-2 x
0.9-1 mm, non-glandular trichomes scattered,
divaricate, 4-6-celled up to 0.7 mm long,
glandular trichomes absent, sessile glands
scattered; lower lobe broadly ovate, 4-4.5 x

4.2- 4.8 mm, non-glandular trichomes sparse,
divaricate, 6-celled up to 1 mm long, glandular
trichomes absent, sessile glands scattered;
filaments filiform, 7-8 x c. 0.2 mm, lilac,
fused for 4-4.5 mm from base; anthers c. 0.5
x 0.4 mm; style filiform, 7-8 mm long, lilac,
bifid for c. 0.5 mm. Fruit calyces 3.5-4 mm
long; upper lobe oblong-ovate, 1.2-2 x 1.3-1.5
mm; lateral lobes lanceolate-falcate, 1.2-2 x
0.7-0.8 mm; lower lobes lanceolate-falcate,

1.2- 1.6 x 0.5-0.6 mm. Nutlets ± circular in
outline, ± flattened, 0.8-0.9 mm long, 0.9-1
mm wide, 0.5-0.6 mm thick, glossy brown,
surface smooth. Fig. 7.

Additional specimens examined : Queensland. Cook
District: 1.8 km W of Mosquito Point Just E of Hunter
Inlet, Temple Bay, Lockhart River Aboriginal Reserve,
May 1992, Fell DF2575 (BRI); Restoration Island, Apr
1995, Le Cussan 285 (BRI); South Pap, Tozers Gap, Jul
1991, Forster PIF9072 (BRI); Puff de Looney Ridge
[Mt Tozer, Iron Range N.P], Jul 1972, Irvine 264 (BRI,
QRS); Brown’s Creek, Pascoe River, Jul 1948, Brass
19590 (BRI); 19 km from the Peninsula Development
road on a track to Wolverton via the Cook tin mine, Jun
1993, Clarkson 10114 & Neldner (BRI).

402

Distribution and habitat : Plectranthus
venustus occurs on Cape York Peninsula
in far north-eastern Queensland in an area
bounded by Mosquito Point in the north to
‘Wolverton’ in the south. Plants occur on
granite pavements and outcrops, but never on
granite rock piles where it is replaced by the
much taller growing Plectranthus excelsus
PI.Forst.

Notes : At Iron Range both Plectranthus
excelsus and P. venustus may occur in close
geographical proximity but have not been
observed growing sympatrically.

Plectranthus venustus appears to have
morphological similarities to P. gratus (Forster
1996) from the Wet Tropics, Queensland.
Observed morphological differences between
these three species are summarized in Table
3.

Austrobaileya 8 ( 3 ): 387-404 ( 2011 )

Map 3. Distribution of Plectranthus venustus in far
north Queensland. Shaded area on map indicates
conservation reserves (National Parks, Forest Reserves
and Conservation Parks).

Table 3. Comparison of some morphological character states between Plectranthus gratus
and P. venustus

Character state

P. gratus

P. venustus

stem indumentum

non-glandular trichomes
antrorse

non-glandular trichomes
divaricate

petiole indumentum

non-glandular trichomes
antrorse, glandular trichomes
sparse, sessile glands absent

non-glandular trichomes
divaricate, glandular
trichomes absent, sessile
glands scattered to dense

laminae form and
colour

ovate; bluish-green above,
silver-green below

lanceolate to narrow-ovate;
silver-green above, silver
below

number of marginal
teeth per side of leaf
lamina

13-15

22-33

pedicels

2.5-3 x c. 0.2 mm; antrorse non-
glandular trichomes

0.5-1 x 0.2-0.3 mm;
divaricate non-glandular
trichomes

inflorescence bracts

not forming a coma; ovate,

1.3-1.8 x 1.3-1.8 mm

usually forming a coma;
obovate to broadly ovate, 2-3
x 2.4-3.2 mm

Conservation status : This species is known
from six localities (two in Iron Range N.P
and one in Restoration Island N.P.) and is not
considered to be threatened at this time. It is

likely that further populations exist within the
area of occurrence which is poorly explored
from a botanical view.

Forster, five new species of Plectranthus

403

Fig. 7. Plectranthus venustus. A. flowering stem x0.6. B. abaxial view of leaf xl. C. adaxial view of leaf xl. D. portion
of inflorescence with two verticillasters, buds and flowers x3. E. lateral view of flower x6. F. face view of flower x6. All
from Forster PIF5388 & Tucker (BR1). Del. W.Smith.

404

Etymology : The specific epithet is based
on the Latin word venustus (beautiful) and
alludes to the form of this plant.

Acknowledgements

The assistance of R. Booth, G. Leiper,
M.T. Mathieson, G. McDonald, K. R. McDonald,
I.Menkins, G. &N. Sankowsky, E. J.Thompson
and M.C.Tucker with new discoveries and
collections of both herbarium and live material
of Plectranthus is gratefully acknowledged.
W. Smith (BRI) provided the illustrations.
Translation of the diagnoses into Latin were
undertaken by P. Bostock (BRI).

References

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- (2010). Lamiaceae. In P.D. Bostock & A.E.

Holland (eds.). Census of the Queensland
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Blake, S.T. (1971). A revision of Plectranthus (Labiatae)
in Australasia. Contributions from the
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Crisp, M.D., Laffan, S., Linder, H.P & Monro, A. (2001).
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Queensland Plants: names and distribution , pp.
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- (1997b). Plectranthus amoenus and P.

thallassoscopicus (Lamiaceae), new species
from north-eastern Queensland, Australia.
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- (1999). Plectranthus fasciculatus (Lamiaceae),

a new species from north-eastern Queensland,
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Names and Distribution of Queensland Plants,
Algae and Lichens , pp. 94-98. Queensland
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(Lamiaceae), a new species from north-east
Queensland. Austrobaileya 7: 707-710.

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inselberg vegetation. In S. Porembski & W.
Barthlott (eds.), Inselbergs. Ecological Studies
146: 391-407.

Iucn (2001). Red List Categories and Criteria: Version
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Wilkiea kaarruana Zich & A.J.Ford (Monimiaceae),
a new species from north-east Queensland

F.Zich 1 & A.J.Ford 2

Summary

Zich, F. & Ford, A.J. (2011). Wilkiea kaarruana Zich & A.J.Ford (Monimiaceae), a new species from
north-east Queensland. Austrobaileya 8(3): 405-411. Wilkiea kaarruana Zich & A.J.Ford endemic
to the Wet Tropics of north-east Queensland is described, illustrated and distinguished from related
Australian species. Notes on habitat, distribution, and conservation status are provided.

Key Words: Monimiaceae, Wilkiea , Wilkiea kaarruana , Australia flora, Queensland flora, new
species, taxonomy, identification key, hyperstigma

'F.Zich, Australian Tropical Herbarium, Sir Robert Norman Building, James Cook University Cairns
Campus, PO Box 6811, Cairns, Queensland 4870, Australia

2 A.Ford, CSIRO, Ecosystem Sciences, Tropical Forest Research Centre, P.O. Box 780, Atherton,
Queensland 4883, Australia

Introduction

Wilkiea F.Muell. (Monimiaceae) is a genus
of approximately 12 species distributed from
Papua New Guinea to Australia (Philipson
1980; Whiffin & Foreman 2007). In their
treatment of the genus for the Flora of
Australia , Whiffin & Foreman (2007) elected
not to describe two poorly known taxa from
north Queensland. Recent collections of
one of these taxa, Wilkiea sp. A (Whiffin &
Foreman 2007) by the authors have provided
for the first time female flowers, and abundant
new male flowering specimens, which have
enabled confident placement of the taxon in
the genus Wilkiea.

The flowers of Wilkiea have been recorded
as possessing a hyperstigma (Endress 1979,
1980); that is, a sticky surface receiving pollen
that is not the stigma itself Mucilaginous
secretion from the stigmas as a reward for
pollinators has been reported for at least one
other genus of Monimiaceae, namely the
Madagascan Tambourissa Sonn. (Lorence
1985), which is pollinated by flies and beetles,
while the pollination of Australian Wilkiea
huegeliana (Tul.) A.DC. appears to be mainly
by thrips (Williams et al. 2001).

Staedler & Endress (2009) have revealed
that all taxa in Monimiaceae that possess a
hyperstigma also have a complex, whorled
floral phyllotaxis, in which merism may
increase or decrease once or several times
between whorls. They call organs tepals when
they are anywhere on the floral cup (regardless
of whether on top of the cup or toward the base
of the cup), and bracts when they are below
the floral cup on the pedicel.

Materials and methods

The study is based upon the examination
of herbarium material from BRI and
CNS (formerly QRS) combined with field
observations. All specimens cited have been
seen by one or both authors.

Measurements of the floral parts of Wilkiea
kaarruana are based on fresh material and
material preserved in 70% ethanol. Common
abbreviations in the specimen citations are:
L.A. (Logging Area), N.R/R. (National Park/
Reserve), S.F./R. (State Forest/Reserve) and
T.R. (Timber Reserve).

NCA is an acronym for the Queensland
Nature Conservation Act (1992) and its
associated schedules. Discussions of
conservation status are made in reference to
the criteria of the IUCN (2001).

Accepted for publication 29 June 2011

406

The abbreviation RE in the distribution and
habitat notes refers to Regional Ecosystem,
descriptions of which can be viewed at (in
this case):

http://www.derm.qld.gov.au/wildlife-ecosystems/
biodiversity/regional_ecosy stems/search.
php?&page=31

Extent of occurrence estimates were
derived from the validation of original
collection localities. These data points were
loaded into ESRI ArcView 3.2 and the draw
polygon feature was used to calculate the area
between the points. The area of occupation
estimates were principally derived from a
digital Regional Ecosystem map supplemented
by expert opinion of vegetation types and
habitats within the Wet Tropics bioregion
(hereafter referred to as the Wet Tropics)
(Environment Australia 2005).

Taxonomy

Wilkiea kaarruana Zich & A. J.Ford, species
nova affinis Wilkiea smithii Whiffin a qua
cortice suberoso, ramulis glabris, pedicellis
glabris, floribus femineis longioribus (22-24
mm longis), et stigmate conico differt. Typus:
Queensland. Cook District: Wooroonooran
National Park, 9.5 km along Maple Creek
Road from South Johnstone River bridge, 20
November 2007, A.Ford 5205 (holo: BRI; iso:
A, BO, CANB, CNS, H, K, L, LAE, MO, NSW,
SYD, MEL, SING, US, Z distribuendi).

Wilkiea sp. Palmerston (B.P.Hyland 80);
Jessup (2007, 2010).

Wilkiea sp. Palmerston (BH 80RFK); Hyland
etal. (2003).

Wilkiea sp. Palmerston; Cooper & Cooper
(2004: 320).

Wilkiea sp. A; Whiffin & Foreman (2007:
81).

Illustration : Cooper & Cooper (2004: 320),
as Wilkiea sp. (Palmerston).

Shrub or small tree 5-12 m tall, androdioecious
or monoecious; bark corky, thick and
longitudinally fissured. Leaves glabrous,
slaty green above when fresh; petiole 5-12
mm long, pink-red; lamina elliptic to oblong-

Austrobaileya 8(3): 405-411 (2011)

elliptic, 6-15 cm long, 2.5-6 cm wide, cuneate
to attenuate at base, entire, acuminate at apex,
mucronate; midrib raised above and below.
Male inflorescence axillary, terminal and
ramiflorous, often fasciculate, 1-4 cm long;
occasionally solitary, mostly triads (botryoids
sensu Endress [2010]), sometimes fascicles
of triads (botryoids), in total 1-15 flowers;
pedicels 5-18 mm long, glabrous; bracts
lanceolate, c. 1.1 mm long. Male flowers
globose to depressed globose, c. 2-3.5 mm
wide, yellow to yellow-orange; tepals rounded
to truncate 4-6, c. 1.8 mm long, glabrous on
both surfaces, two pairs around ostiole, third
pair if present near base of floral cup; stamens
4, c. 1.1 mm long, in equal pairs, outer 2
positioned on floral cup wall and higher than
inner two which are positioned on the base
of receptacle, anthers opening by apical slit
curving over apex, filaments very short, hairy.
Female inflorescence terminal, 3-5 cm long,
fascicles of triads (botyroids), in total 6-14
flowers, pedicels 11-24 mm long, glabrous.
Female flowers depressed globose, c. 7 mm
diameter, yellow-orange fading to green at
base; tepals 4-6, 2 pairs around ostiole, outer
pair thin, inner pair greatly thickened, forming
a floral cup, third pair when present often
consisting of one tepal near the base of floral
cup and the opposing tepal on the side of the
floral cup; carpels 21-26, angular, tapering,
densely pubescent with golden, more or less
appressed hairs; stigma short, conical and
black; staminodes absent. Fruiting receptacle,
6—10(—15) mm wide, pubescent; mature fruit
not known. Fig. 1.

Additional selected specimens examined: Queensland.
Cook District: Big Tableland via Cooktown, s.dat, Jack
inEldridge s.n. (BRI [AQ63833]); Gap Creek, c. 22 miles
[36 km] S by E of Cooktown, Oct 1969, Smith s.n. (BRI
[AQ339983]); loc. cit., Smith s.n. (BRI [AQ339984]); loc.
cit., Smith s.n. (BRI [AQ339988]); Daintree N.R, Cedar
Bay Section, Gap Creek, Nov 2000, Forster PIF26462
et al. (BRI); Daintree N.R, Daintree River headwaters.
May 1998, Forster PIF22947 et al. (BRI); Mitchell
Bomber crash site. East Bartle Frere, Wooroonooran
N.R, Nov 1994, Hunter 2147 (BRI); Jordan L.A., 16.5
km SE of Millaa Millaa, site 2, Oct 1988, Jessup GJM
2095 et al. (BRI); Wooroonooran N.R, 1.1 km along K-
Tree road from Kaarru Creek crossing, Nov 2007, Ford
5208 1 (BRI, CNS); Wooroonooran N.R, 1.1 km along K-
Tree road from Kaarru Creek crossing, Nov 2008, Zich
626 x (BRI, CNS); Wooroonooran N.R, Maple Creek
Road, CSIRO Climate Change Plot, Charappa, Nov

Zich & Ford, Wilkiea kaarruana (Monimiaceae)

2008, Zich 62? (BRI, CANB, CNS); Palmerston F.R.,
old logging road towards Karang-Garee Falls, off South
Johnstone Forestry road, Oct 2003, Ford 4174 & Holmes
(BRI, CNS); ( - flowering specimen).

Distribution and habitat : Wilkiea kaarruana
is endemic to the Wet Tropics bioregion
in north-eastern Queensland, where it is
currently known to occur from the Cooktown
area (Big Tableland and Mt Finnigan environs)
to the South Johnstone River (south-east of
Millaa Millaa) (Map 1). However, within this
large range Wilkiea kaarruana is confined to
four distinct and disjunct populations (from
north to south):

1. Mt Finnigan area (from Big Tableland to
Gap Creek)

2. Daintree River headwaters

3. Mt Bartle Frere

4. South Johnstone River catchment.

It inhabits predominantly the wetter and
more mountainous notophyll vine-forests/
rainforests on soils derived from granite, fine
grained metasediments (including mudstone)
and basalt. However, it is more commonly
encountered on granitic substrates, with
occurrences on basalt and metasediments
being less common. Common canopy
species that grow in association with Wilkiea
kaarruana include: Cardwellia sublimis
F.Muell., Carnarvonia araliifolia F.Muell.,
Cryptocarya mackinnoniana F.Muell.,
Doryphora aromatica (F.M.Bailey) L.S.Sm.,
Elaeocarpus ruminatus F.Muell., Flindersia
bourjotiana F.Muell., Flindersia pimenteliana
F.Muell., Franciscodendron laurifolium
(F.Muell.) B.Hyland & Steenis, Sloanea
macbrydei F.Muell., Syzygium johnsonii
(F.Muell.) B.Hyland, Syzygium papyraceum
B. Hyland and Xanthophyllum octandrum
(F.Muell.) Domin. Common small trees and
shrubs throughout most of its range include:
Apodytes brachystylis F.Muell., Ardisia
brevipedata F.Muell., Bobea myrtoides
(F.Muell.) Valeton, Bubbia semecarpoides
(F.Muell.) B.L.Burtt, Chionanthus axillaris
R.Br., Citronella smythii (F.Muell.)
R.A. Howard, Gossia spp., Pilidiostigma
tropicum L.S.Sm., Pittosporum rubiginosum
A.Cunn., Polyscias australiana (F.Muell.)

407

Philipson and Psychotria spp. Altitudinal
range, from existing specimens, is 60-1090
m.

Wilkiea kaarruana has been collected
or reliably reported in the following REs:
7.8.2a (rarely); 7.11.1a (occasionally); 7.12.1a
(occasionally), 7.12.16a (commonly) and
7.12.19a (rarely).

Phenology : Flowers have been recorded in
October and November; immature fruits have
been recorded in March.

Affinities: Wilkiea kaarruana is

morphologically most similar to W. smithii
and W. hylandii Whiffin. However these
species are easily distinguished by several
key features and geographic distribution.
All three species have pubescent carpels and
glabrous to glabrescent adaxial leaf surfaces.
A comparison of diagnostic differences
between these three species (and Wilkiea
austroqueenslandica ) is provided in Table 1.

Whiffin (2007) suggests that Wilkiea
austroqueenslandica, W. hylandii and
W. smithii form a “distinctive group”, as
they all have pubescent inflorescences and
carpels. Although W. kaarruana has glabrous
inflorescences, the carpels are distinctly
pubescent. Molecular studies by Renner et
al. (2010, Fig. 2) and Renner pers. comm.
(2009) place Wilkiea kaarruana close to
W. austroqueenslandica Domin., W. smithii
and W. hylandii.

Notes : In habitat Wilkiea kaarruana is easily
recognized by the thick and longitudinally
fissured pale corky bark. A specimen from the
Big Tableland ( Jack in Eldridge s.n .) consists
of a piece of bark 20mm thick mounted on a
sheet and the name “Corkwood Tree” on the
original label. No other species of Wilkiea , or
Monimiaceae, in Australia have this striking
feature though it has been reported for at least
two species of Monimiaceae on the island of
New Guinea, currently assigned to the island
endemic genus Kairoa Philipson (Renner &
Takeuchi 2009). Recent molecular research
on Kairoa indicates it should be included with
Wilkiea , along with Kibara Endl., which is
likewise embedded among species of Wilkiea
(Renner et al. 2010).

408

Austrobaileya 8(3): 405-411 (2011)

Table 1. Morphological comparison between Wilkiea austroqueenslandica , W. hylandii ,

W. kaarruana and W. smithii

Character

state

W. austroqueenslandica

W. hylandii

W. kaarruana

W. smithii

Bark

finely wrinkled

not recorded (but
not corky)

corky

tessellated

flaky

Leaf

indumentum

glabrous

pubescent when
young, glabrescent
or persisting on
midrib

glabrous

densely
pubescent
when young,
esp. midrib,
glabrescent

Leaf margin

prominently, irregularly
toothed

prominently
toothed in upper
half

entire

entire

Male

inflorescence

position

axillary

axillary,

sometimes

fasciculate

axillary,
terminal and
ramiflorous,
often fasciculate

axillary,

sometimes

fasciculate

Male

inflorescence

triad or fascicle of triads

triad or fascicle of
triads

occasionally
solitary, mostly
2-flowered or
sometimes triads
or fascicle of
triads

triad or
fascicle of
triads

Male flower

c. 8 mm

4-5 mm

5-18 mm

6-8 mm

pedicel
length &
indumentum

densely pubescent

densely pubescent

glabrous

densely

pubescent

Male flowers
tepal number

8 as 4 pairs

6, with 2 pairs
around ostiole,
and 1 lower on
floral cup

4 to 6. 2 as
pairs around an
ostiole. 3rd pair,
if present at base
of floral cup

8 in 2 whorls

Male flowers

stamen

number

30

(6)8(10)

4

usually 4

Female

inflorescence

position

axillary

axillary or
terminal

terminal

axillary or
terminal

Female

10-15 mm

7-15 mm

22-24 mm

5-8 mm

flower pedicel
length &
indumentum

densely silky pubescent

densely pubescent

glabrous

densely

pubescent

Female flower
tepal number

4

4

4 or 6

4

Carpel

number

35

40

21-26

20-40

Stigma shape

hemispherical

hemispherical

conical

hemispherical

Zich & Ford, Wilkiea kaarruana (Monimiaceae)

Although Wilkiea kaarruana is a small
tree, it is remarkable that there are so few
fertile specimens in herbaria. The few
flowering collections have sometimes made
it difficult to determine levels of dioecy and
monoecy. As noted by Whiffin & Foreman
(2007) the genus Wilkiea is monoecious or
perhaps rarely dioecious, and “It is likely
that many (perhaps all) species described
as dioecious have male and female flowers
on the same plant but usually not at the
same time. For example, many plants with
mature fruit have young flowers that are
male”. Observations made during this study
corroborate these statements, with Ford 5208
having mature female flowers and younger
male flowers. These male flowers would
likely come into anthesis after the female
flowers were fertilized. Also, despite regular
monitoring of several plants before and after
flowering, we found no evidence of male plants
(Ford 5205 and Zich 628 ) ever producing
female flowers. Accordingly, we suggest that
Wilkiea kaarruana is androdioecious (male
dioecious) and monoecious (having pure male
plants growing together with monoecious
plants). Further field observations are
required to assess the reproductive strategy
within Wilkiea kaarruana , and indeed for all
Australian Monimiaceae.

Floral observations of female Wilkiea
kaarruana conducted as part of this study
have confirmed the presence of a hyperstigma.
The outline of the flower is globular, the inner
pair of tepals is greatly thickened and appears
to produce mucilage at anthesis. The amount
of mucilage in dissected flowers varied but
was observed to fill the floral cup creating a
continuous cover over the carpels to the apical
pore (see Fig. 1). Despite examination of many
flowers no insects or their larvae were found,
though possible evidence of insect damage
was observed in the partial destruction of the
apical pore as described and illustrated by
Endress (1980: 85, fig.35).

The presence of a third whorl of tepals in
both male and female flowers in W. kaarruana
is not unusual within Wilkiea ; however, it is
more usual for male flowers to possess more
whorls than female flowers (Whiffin 2007).

409

Leaf oils in Australian species of
Wilkiea have been studied by Brophy et
al. (2009). Wilkiea kaarruana produced a
sesquiterpenoid oil similar to the related
Wilkiea smithii Whiffin; although this
latter species also contained very small
amounts of monoterpenes. Wilkiea
austroqueenslandica Domin, another
closely related species, yielded an oil also
dominated by sesquiterpenes, but had a
much higher proportion of monoterpenes.
Wilkiea hylandii Whiffin was not sampled
in their study.

Conservation status : Most existing collections
of this species have been made within the
World Heritage Area of the Wet Tropics.
Wilkiea kaarruana has been collected in
Daintree and Wooroonooran National Parks.
The species has a wide, albeit restricted and
disjunct geographical range with an area of
occupation estimated to be 110 km 2 . It is not
considered at risk.

Etymology : The specific epithet honours
Kaarru (pronounced Kaa-roo) Creek and
Kaarru Logging Area, where several
collections of the species, including the first
female flowering collection were made. Now
known as Wooroonooran National Park, the
area was previously referred to as State Forest
Reserve 756.

Acknowledgements

The authors wish to thank Will Smith
for the illustrations. Peter Bostock provided
the distribution map and Lyn Craven the
translation of the diagnosis into Latin. Peter
Endress and Susanne Renner improved an
earlier draft. Permits to collect in the Wet
Tropics were issued by the Department of
Environment & Resource Management. The
curators and staff at BRI are thanked for
allowing access to specimens and the use of
their facilities.

410

Austrobaileya 8(3): 405-411 (2011)

Fig. 1. Wilkiea kaarruana. A. large branch showing longitudinal corky fissures x 1. B. transverse section of large
branch x 2. C. branchlet with male inflorescences x 0.5. D. leaf adaxial surface with extra venation detail x l.E. lateral
view of male flower showing off-set tepal on floral tube and bract on pedicel x 10. F. face view of male flower showing
tepals around ostiole and anthers x 12. G. lateral view of female flower showing off-set tepals on floral cup x 10. H.
face view of female flower showing tepals around ostiole and off-set tepal on floral cup x 12.1. section of female flower
showing hairy carpels and mucilaginous cover from the hyperstigma x 12. A-B from Forster PIF26462 et al. (BRI);
C-F from FordAF5205 (CNS); G-I from FordAF5208 (CNS). Del. W.Smith

Zich & Ford, Wilkiea kaarruana (Monimiaceae)

References

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(1965). Flora of Java, Vol. 2. N.V.P. Noordhoff:
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Brophy, J.J., Goldsack, R.J. & Forster, RI. (2009).
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oils of the Australian species of Endressia ,
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Cooper, W. & Cooper, W.T. (2004). Fruits of the
Australian Tropical Rainforest. Nokomis
Editions: Melbourne.

Endress, P.K. (1979). Noncarpellary pollination and
‘hyperstigma’ in an angiosperm ( Tambourissa
religiosa, Monimiaceae). Experientia 35: 45.

- (1980). Ontogeny, function and evolution of

extreme floral construction in Monimiaceae.
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- (2010). Disentangling confusions in

inflorescence morphology: Patterns and
diversity of reproductive shoot ramification
in angiosperms. Journal of Systematics and
Evolution 48(4): 225-239.

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Biogeographic Regionalisation for Australia
(IBRA) and Development of Version 5.1
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Version 6.1 (Digital Data, metadata), [cited 24
June 2009], http://www.environment.gov.au/
parks/nrs/science/bioregion-framework/ibra

Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray,
B. & Elick, R.W. (2003). Australian Tropical
Rain Forest Plants. Trees, Shrubs and Vines.
CD-ROM. CSIRO Publishing: Melbourne.

Jessup, L.W. (2007). Monimiaceae. In P.D.Bostock &
A.E.Holland (eds.). Census of the Queensland
Flora 2007 ’ p.l 16. Environmental Protection
Agency: Brisbane.

- (2010). Monimiaceae. In P.D.Bostock &

A.E.Holland (eds.). Census of the Queensland
Flora 2010 , p. 111. Queensland Herbarium,
Department of Environment & Resource
Management: Brisbane.

Lorence, D.H. (1985). A monograph of the Monimiaceae
(Laurales) in the Malagasy Region (Southwest
Indian Ocean). Annals of the Missouri Botanical
Garden 72: 1-165.

Philipson, W.R. (1980). The genus Wilkiea (Monimiaceae)
in Papua New Guinea. Blumea 26: 365-366.

Renner, S.S. & Feil, J.P (1993). Pollinators of tropical
dioecious angiosperms. American Journal of
Botany 80: 1100-1107.

411

Renner, S.S., Strijk, J.S., Strasberg, D. & Thebaud,
C. (2010). Biogeography of the Monimiaceae
(Laurales): a role for East Gondwana and long¬
distance dispersal, but not West Gondwana.
Journal of Biogeography 37: 1227-1238.

Renner, S.S. & Takeuchi, W.N. (2009). A phylogeny
and revised circumscription for Kairoa
(Monimiaceae), with the description of a new
species from Papua New Guinea. Harvard
Papers in Botany 14: 71-81.

Sampson, F.B. (1969). Studies on the Monimiaceae.
I. Floral morphology and gametophyte
development of Hedycarya arborea J.R. et G.
Forst. (subfamily Monimioideae). Australian
Journal of Botany 17: 403-424.

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lability of floral phyllotaxis in the Pluricarpellate
families of core Laurales (Gomortegaceae,
Atherospermataceae, Siparunaceae,

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Sciences 170: 522-550.

Whiffin, T. & Foreman, D.B. (2007). Monimiaceae.
Flora of Australia 2: 65-91. ABRS/CSIRO
Publishing: Melbourne.

Williams, G.A., Adam, P. & Mound, L.A. (2001). Thrips
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1 - 21 .

Map 1. Distribution of Wilkiea kaarruana in Queensland.
Shaded areas indicate conservation reserves.

New and reinstated species of the Solanum
ellipticum R.Br. (Solanaceae) species group

A.R.Bean

Summary

Bean, A.R. (2011). New and reinstated species of the Solanum ellipticum R.Br. (Solanaceae) species
group. Austrobaileya 8(3): 412-430. The Solanum ellipticum species group is endemic to Australia,
and is here considered to comprise 13 species. Solanum lithophilum F.Muell. and S. cleistogamum
Symon are reinstated. Solanum aridicola A.R.Bean, S. callosum A.R.Bean, S. emmottii A.R.Bean
and S. unispinum A.R.Bean are described as new. The four new species and Solanum lithophilum are
illustrated and distribution maps are provided for them. A key to the identification of species in the
Solanum ellipticum group is presented.

Key Words: Solanaceae, Solanum, Solanum aridicola, Solanum callosum, Solanum ellipticum,
Solanum emmottii, Solanum lithophilum, Solanum unispinum, Australia flora. New South Wales
flora. Northern Territory flora, Queensland flora. Western Australia flora, taxonomy, new species,
identification key

A.R. Bean, Queensland Herbarium, Department of Environment & Resource Management, Brisbane
Botanic Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia. E-mail: tony.bean@
derm.qld.gov.au

Introduction

Solanum ellipticum was named by Robert
Brown in 1810, based on type material
collected near St Lawrence on the central
Queensland coast. For many years, the name
has been widely applied to Solanum species
with the following characteristics: prostrate or
low growing, stems moderately prickly, calyx
prickly; leaves ovate to elliptical with entire
(or repand) margins, densely stellate-hairy
at least on the lower surface; inflorescences
racemose; fruits green, yellow-green or
almost white at maturity; calyx not accrescent.
Solanum specimens matching these criteria
occur in all mainland states except Victoria.

The current author was able to distinguish
some taxa (, Solanum crebrispinum A.R.Bean,
S. senticosum A.R.Bean) within S. ellipticum
sens. lat. based on herbarium material (Bean
2004). Subsequent field studies have provided
new insights and revealed characters that
are not readily apparent from the standard
herbarium specimen. For instance, there
are consistent differences between taxa in
growth habit (Figs. 1 & 2), corolla shape
and indumentum, style orientation (erect or
eccentric), stigma size and colour, fruit colour
and placentation, and pericarp thickness.

Accepted for publication 27 June 2011

Bean (2004) included 10 species within
the informal Solanum ellipticum group
(Group 27), which is endemic to Australia.
The membership of that group needs
alteration. On morphological grounds, the
author now recognises that the affinities
of S. crassitomentosum Domin lie with
S. furfuraceum R.Br. (Group 27B), that
S. angustum Domin belongs near A graniticum
A.R.Bean in Group 25, and that A argopetalum
A.R.Bean belongs in the S. esuriale Lindl.
group (Group 27C). Additions to the group
comprise the reinstated S. cleistogamum
Symon and S. lithophilum F.Muell., and the
four species named in this paper (S. unispinum,
S. emmottii, S. callosum and S. aridicola).
With these changes, Group 27 now comprises
13 species, namely S. aridicola, S. callosum,
S. cleistogamum, S. crebrispinum A.R.Bean,
S. dianthophorum Dunal, S. ellipticum,
S. emmottii, S. horridum Dunal ex Poir.,
S. lithophilum, S. quadriloculatum F.Muell.,
S. senticosum A.R.Bean, S. terraneum Symon
and S. unispinum.

Solanum terraneum is tentatively retained
in the S. ellipticum group following Symon
(1981), but no specimens have been seen by
the present author, and so it is not included
in the key to species and is not considered
further in this paper.

Fig. 1. Plant of Solanum ellipticum at Homevale National Park ( Bean 30650 et ah). Photograph: A.R.Bean

Full descriptions are given for the new
species, the reinstated species {Solanum
cleistogamum and S. lithophilum ), and for the
recircumscribed Solanum ellipticum.

Materials and methods

Relevant specimens have been examined in
the herbaria BRI, MEL, NSW and NT; as have
specimens of Solanum ellipticum sens. lat.

from CANB, and material of S. cleistogamum
from PERTH.

Measurement techniques and terminology
are as per Bean (2004). All species except
Solanum dianthophorum and S. terraneum
have been examined in the field. Species
treatments are in alphabetical order.

414

Austrobaileya 8(3): 412^130 (2011)

Taxonomy

Key to the species in the Solanum ellipticum group

1 Stems and branchlets without prickles.S. dianthophorum

1. Stems and branchlets sparsely to densely prickly.2

2 Style protruding between anthers or slightly bent, displacing one anther,

erect for most of its length but recurved near apex.3

2. Style bent strongly near base, displaced from the centre of the flower

(eccentric).10

3 Stems ultimately ascending or erect; bushy plants.4

3. Stems ultimately prostrate or procumbent; ground-covering plants.8

4 Calyx prickles 40-70, very stout and pungent.5

4. Calyx prickles rather flimsy, 1-40, or sometimes absent.6

5 Leaves 1.6-1.8 times longer than broad, base obtuse or cordate; ovary

and style with very shortly stalked glandular hairs; calyx stellate hairs

0.7-0.9 mm across.S. crebrispinum

5. Leaves 1.9-2.7 times longer than broad, base cuneate; ovary and style

glabrous; calyx stellate hairs 0.4-0.6 mm across.S. senticosum

6 Leaves 2.2-3.5 cm long; calyx prickles 0-5; inflorescences 1 or 2

flowered.S. unispinum

6. Leaves 4.9-14.5 cm long; calyx prickles 5-40; inflorescences 7-15 flowered

7 Stellate hairs consistently present on inner surface of corolla; branchlets

terete; prickles glabrous.S. callosum

7. Stellate hairs consistently absent from inner surface of corolla; branchlets

usually ridged; prickles with stellate hairs attached at their bases

.S. quadriloculatum

8 Calyx prickles >50; fruits 18-22 mm diameter; inflorescences 1(—2)-

flowered.S. horridum

8. Calyx prickles 5-50; fruits 9-17 mm diameter; inflorescences 1-6-

flowered.9

9 Fruits 9-12 mm diameter; anthers 2-2.8 mm long; central ray of stellate

hairs 1-2 times as long as laterals.S. cleistogamum

9. Fruits 13-17 mm diameter; anthers 4-5.5 mm long; central ray of stellate

hairs 0.5-1 times as long as laterals.S. ellipticum

10 Plant prostrate to procumbent; calyx prickles pungent, 3.5-8 mm long

.S. aridicola

10. Plant bushy, with ascending stems; calyx prickles flimsy, 0.5-4 mm

long.11

11 Branchlet hairs not floccose (stalks 0-0.2 mm); calyx prickles 0.5-2 mm

long; corolla 9-13 mm long.S. emmottii

11. Branchlet hairs floccose (stalks 0-2 mm); calyx prickles 2-4 mm long;

corolla 14-20 mm long.S. lithophilum

Bean, Solarium ellipticum group

415

Fig. 2. Population of Solarium lithophilum , north of Alice Springs (site of collection for Bean 25457 & Albrecht).
Photograph: A.R.Bean

Solanum aridicola A.R.Bean species nova

S. horrido affinis sed ab eo inflorescentiis
3-8-floris, praesentia pilorum stellatorum
longe stipitatorum in pagina superiore
foliorum et marginibus foliorum aequabiliter
integris differens. Typus: Northern Territory.
Larapinta trail, Standley Chasm area. c.
40 km W of Alice Springs, 1 August 2006,
A.R.Bean 25449 & D.E.Albrecht (holo: BRI;
iso: CANB, MEL, NSW, NT).

S. ellipticum form “b”; Symon (1981).

S. ellipticum var. Ranges (D.E.Albrecht
5807); Kerrigan & Albrecht (2007), Albrecht
et al. (2007: 176).

Herbaceous resprouter or rhizomatous
perennial shrub 0.1—0.3 m high. Stems initially
ascending, then prostrate or sprawling. Adult
branchlets yellow or rusty; prickles 35-56
per cm, straight, acicular, 4-12 mm long,

14-20 times longer than wide, glabrous;
stellate hairs very dense, lateral rays 8-9,
porrect or ascending, central ray 1-2 times
as long as laterals; Type 2 hairs absent. Adult
leaves elliptical or ovate, entire or shallowly
lobed throughout with 2-3 on each side;
lobes obtuse, lobing index 1-1.1; lamina
2.8-9.8 cm long, 1.3-4.4 cm wide, 2.1-2.8
times longer than broad, apex acute, base
cuneate or obtuse, oblique part 0-4.5 mm
long, obliqueness index 0-6%; petioles 1.3-4
cm long, 26-56% length of lamina, prickles
present. Upper leaf surface green; prickles
present on midvein only or present on midvein
and lateral veins, 1-20, straight, acicular,
3.5-13 mm long; stellate hairs distributed
throughout, very dense; lateral rays 8, porrect
or ascending, central ray 1-2 times as long
as laterals; simple hairs absent; Type 2 hairs
absent. Lower leaf surface yellowish; prickles
present on midvein only or present on midvein

416

and lateral veins, 3-35, straight, acicular;
stellate hairs very dense, lateral rays 8-12,
porrect, ascending or multiradiate, central ray
0.9-2 times as long as laterals; simple hairs
absent; Type 2 hairs absent. Inflorescence
supra-axillary, cymose (pseudo-racemose),
common peduncle 11-27 mm long, rachis
prickles present, 2-8-flowered. Flowers 5-
merous. Pedicels at anthesis 3-5 mm long,
same thickness throughout, prickles absent
or present; calyx tube 2-3 mm long; lobes
attenuate, 3.5-8 mm long; prickles 22-50
per flower, 3.5-8 mm long; stellate hairs very
dense, lateral rays 7-12, central ray 1-2 times
as long as laterals; simple hairs absent; Type
2 hairs absent. Corolla rotate, 9-15 mm long,
purple, inner surface glabrous; anthers 3.6-
4.3 mm long; filaments c. 1 mm long; ovary
glabrous. Functional style 8-8.5 mm long,
strongly bent near base, not touching anthers,
eccentric, glabrous; stigma expanded, darker
than style, obscurely lobed. Fruiting calyx
lobes less than half length of mature fruit.
Mature fruits 1-3 per inflorescence, oblate
or globular, 18-21 mm diameter, brown or
yellow, 2-locular; placenta stalked, anvil¬
shaped, pericarp 0.5-0.7 mm thick; pedicels
7-12 mm long. Seeds 1.8-2.5 mm long, white
or pale yellow. Fig. 3.

Additional selected specimens examined: Northern
Territory. Un-named gorge between Hugh and Spencer
Gorge, Mar 1994, Albrecht 5807 (AD, NT); Valley of
Eagles, 35 miles [58.3 km] ENE of Alice Springs, Oct
1966, Beauglehole 20671 (NT); James Range, 3 miles
[5 km] N of Hugh River, Aug 1967, Beauglehole 24592
(AD, NT); Standley Chasm, MacDonnell Ranges, Jul
1968, Orchard 818 (AD, NT); Palm Valley, Sep 1970,
Late 790 (NT); Kings Canyon, Dec 1968, Latz 321 (NSW,
NT); ‘Derry Downs’, Dulcie Range, Oct 1974, Latz 5828
(NT); Redbank Gorge, 3 Jul 1985, Leach 666 (MEL,
NT); Emily Gap, 8 miles [13 km] E of Alice Springs, Feb
1968, Nelson 1618 (MEL, NSW, NT); Stuart Highway,
10.7 km N of Aileron, Aug 2006, Bean 25455 & Albrecht
(BRI, NT).

Distribution and habitat : Solanum aridicola
is known only from the southern parts of
the Northern Territory (Map 1), although it
probably occurs in adjacent parts of Western
Australia and South Australia. It inhabits
granite or quartzite hills and ranges with
shallow sandy loam soils. Associated species
include Eucalyptus intertexta R.T.Baker,
E. terminalis F.Muell. and Grevillea
wickhamii Meisn.

Austrobaileya 8(3): 412-430 (2011)

Phenology : Flowers have been recorded
for July and August; fruits in August and
February.

Notes : Solanum aridicola is most closely
related to S. horridum as both species are more
or less prostrate, with very prickly stems and
leaves and have flowers and fruits of about
the same size and colour. Solanum aridicola
is most readily distinguished by the 3-8-
flowered inflorescences (1 or 2-flowered in S.
horridum ), and by the eccentric style in the
flower. It also differs from Solanum horridum
by the entire leaves (often shallowly lobed in
S. horridum) and the upper leaf surface with
many long-stalked stellate hairs (all ± sessile
in S. horridum).

Etymology : From the Latin aridus meaning
‘arid’ and -cola meaning ‘to inhabit’. This
refers to the arid landscape that the species
inhabits.

Solanum callosum A.R.Bean species nova

S. quadriloculato affinis sed ab eo fasciculis
pilorum stellatorum in corollam, lobis
calycum apicibus longis linearibus praeditis,
caulibus teretibus et ovario interdum 2-
loculari differens. Typus: Queensland.
Gregory South District: 20.2 km WSW of
Eromanga, on Cooper Development Road, 27
August 2010, A.R. Bean 30027 (holo: BRI [1
sheet + spirit]; iso: NSW).

S. ellipticum R.Br., and. non Bean (2004),
pro parte.

Illustrations : Cunningham et al. (1981: 592),
as S. quadriloculatum ; Brooke & McGarva
(1998: 112), as S. ellipticum.

Rhizomatous perennial shrub 0.15-0.6
m high. Stems initially sprawling, then
spreading to erect. Adult branchlets yellow,
rusty or brown; prickles 8-70 per cm, straight,
acicular, 2-7 mm long, 14—18 times longer
than wide, glabrous; stellate hairs very dense,
0.6-1.1 mm diameter, stalks 0-0.2 mm long,
lateral rays 7-8, porrect or ascending, central
ray 0.7-1.4 times as long as laterals; Type 2
hairs absent. Adult leaves ovate or broadly
ovate, entire, lamina 4.9-9 cm long, 3-4.8
cm wide, 1.5-2.4 times longer than broad,
apex obtuse or acute, base cuneate or obtuse,
oblique part 1-3.5 mm long, obliqueness

Bean, Solanum ellipticum group

417

Fig. 3. Solanum aridicola. A. flowering branchlet * 0.8. B. flower with corolla and calyx lobes removed x 4. C. pedicel,
calyx and style x 4. D. corolla x 2. E. ovary and style x 4. F. cross-section of mature fruit x 2. All from Bean 25449 &
Albrecht (BRI).

index 1-4%; petioles 1.2-3 cm long, 17-33%
length of lamina, prickles present. Upper leaf
surface grey; prickles absent or with 1-4 on
midvein only, straight, acicular, 2-4.5 mm
long; stellate hairs distributed throughout,
dense or very dense; 0.05-0.2 mm apart,
0.5-0.7 mm across, stalks 0-0.15 mm long,
lateral rays 6-8, porrect or ascending, central
ray 0.7-1.3 times as long as laterals; simple
hairs absent; Type 2 hairs absent. Lower leaf

surface white or grey; prickles 3-10, present
on midvein only or present on midvein and
lateral veins, straight, acicular; stellate hairs
very dense, 0.05-0.1 mm apart, 0.6-0.9 mm
diameter, stalks 0-0.3 mm long, lateral rays
7-8, porrect, central ray 0.7-1.3 times as long
as laterals; simple hairs absent; Type 2 hairs
absent. Inflorescence supra-axillary, cymose
(pseudo-racemose), common peduncle 15-
41 mm long, rachis prickles present, 7-11-

418

flowered, with all flowers bisexual. Flowers
5-merous. Pedicels at anthesis 5-11 mm long,
same thickness throughout, prickles absent
or present. Calyx tube at anthesis 2.5-4 mm
long; lobes deflate or attenuate, 1.5-7.5 mm
long; prickles 14-40 per flower, 1-3 mm long;
stellate hairs very dense, white or purplish,
0.4-0.8 mm across, stalks 0.1-0.5 mm long,
lateral rays 7-8, central ray 0.8-1.4 times as
long as laterals; simple hairs absent; Type 2
hairs absent. Corolla shallowly lobed, 9-14
mm long, purple, inner surface densely
stellate-hairy at distal end; anthers 4.4-6.1
mm long; ovary glabrous; style 7.5-8.5 mm
long, protruding between anthers, glabrous;
stigma unexpanded, same colour as style,
obscurely lobed. Fruiting calyx lobes less
than or more than half length of mature fruit,
prickles 2-4 mm long. Mature fruits 2-4 per
inflorescence, globular, 18-20 mm diameter,
green, 2-locular or 4-locular, placenta stalked,
anvil-shaped, interior moist but not juicy,
pericarp 3-3.3 mm thick. Pedicels at fruiting
stage 8-21 mm long. Seeds pale yellow, 2.5-
2.7 mm long. Fig. 4.

Additional specimens examined: Queensland. Mitchell
District: 42.7 km along Silsoe Road, W of Longreach,
May 2004, Bean 22213 (BRI); Longreach, Thompson
River, Apr 1963, Gittins 670 (BRI); Milo Station, Old
Gooyea Outstation ruins, near Powell Creek, NNW of
Adavale, Aug 2009, Forster PIF35347 & Thomas (BRI,
NSW); ‘Springdale’, N of Aramac, Jul 2000, Fensham
3947 (BRf); road from Adavale to Windorah, 64 miles
[102 km] WNW of Adavale, Jun 1967, Gittins 1232
(BRI); Idalia N.P., SW of Blackall, Jun 1990, Rogers s.n.
(BRI [AQ624125]). Gregory North District: Kangaroo
Mountains, between Bedourie and Windorah, Jun 1978,
Althofer 8363 (BRI). Gregory South District: 3.7 km
From Poison Tank Bore, GiGi Paddock, Mt Margaret
Station, 30 km SW of Eromanga, Oct 1995, Stol &
Landsberg QSL382 (CANB); near Jackson, south-west
Qld, Jul 2003, Hines s.n. (BRI [AQ779478]); c. 40 miles
[64 km] W of Windorah, Jun 1949, Everist 3894 (BRI);
19.1 km W of central Tank turnoff, Thargomindah
- Nockatunga Road, Sep 1990, Prendergast HDP346
(BRI). Warrego District: Eulo sports ground, 0.8 km
S of Eulo, Aug 2006, Bean 25634 (BRI); 42.3km from
Charleville, towards Quilpie, Aug 2010, Bean 29925
(BRI, MEL); ‘Gilruth Plains’, E of Cunnamulla, May
1939, Blake 14034 (BRI); Currawinya N.P, 4 km WNW
of Gidgee Bore, Mar 1997, Forster PIF20444 & Watson
(BRI, DNA, MEL); c. 27 km NE of Thargomindah, on
road to Quilpie, Sep 1973, Henderson H2083 & Boyland
(BRI); ‘Yarronvale’, 66 miles [106 km] from Charleville
on Quilpie road, Sep 1963, Everist 7520 (BRI). Maranoa
District: ‘Murra Murra’, River paddock, at Brumby
Lake, SW of Bollon, Feb 2007, Eddie CPE1091 (BRI).

Austrobaileya 8(3): 412-430 (2011)

New South Wales. North Far Western Plains: Ursino -
Yamba Station road, 30 miles [48 km] W of Wanaaring,
Oct 1963, Constable 4583 (BRI, NSW); 14 miles [23 km]
W of Wanaaring, Dec 1969, Martensz 260 (AD, CANB,
NSW); Peery Station, Apr 1963, Bailey 2862 (CANB).
North Western Plains: 65.7 km from Bourke - Cobar
Road, towards Coolabah, Mar 2008, Bean 27637 (BRI);
0.1 km W of Condobolin - Cobar Road, c. 40 km N of
Condobolin, Mar 2008, Bean 27694 (BRI); Coolabah,
Apr 1900, Helms s.n. (NSW).

Distribution and habitat : Solanum callosum
is widespread from north of Muttaburra in
central Queensland to Condobolin in central
New South Wales (Map 2). It grows on plains
and low stony hills on sandy loam to clay
loam soils. Associated tree species include
mulga {Acacia aneura F.Muell. ex Benth.),
A. excelsa Benth., Eucalyptus terminalis
F.Muell. and E. populnea F.Muell.

Phenology : Flowers are recorded for all
months of the year. Fruits are recorded
between February and September.

Notes: This species is most closely allied to
Solanum quadriloculatum ; however, there
a number of differences; S. callosum has a
cluster of stellate hairs on the distal portion
of the inner surface of the corolla, with
scattered stellate hairs extending down the
midvein for about half the corolla length (in
S. quadriloculatum the corolla is glabrous,
except for two or three collections which bear
simple hairs at the distal end of the lobes);
calyx lobes narrowly deflate with long linear
apices (broadly deflate, with or without short
linear apices for S. quadriloculatum ); stems
terete (usually ridged for S. quadriloculatum );
no stellate hairs on prickles (stellate hairs on
at least some prickles for S. quadriloculatum );
ovary 2- or 4-locular (consistently 4-locular
for S. quadriloculatum) ; fruits 18-20 mm
diameter (14-17 mm for S. quadriloculatum).

Etymology: The species epithet is from the
Latin callosus meaning tough, hard, or thick-
skinned. This is given in reference to the
fruits that are hard when dry, and for the very
thick pericarp.

Bean, Solanum ellipticum group

419

Fig. 4. Solanum callosum. A. flowering branchlet x 0.6. B. oblique view of an open flower x 2. C. base of flower
showing calyx and corolla x 2. D. mature fruits attached to rachis x 1.5. E. cross-section of a 4-locular fruit x 2. F.
cross-section of a 2-locular fruit x 2. A from Gittins 670 (BRI); B,C from Bean 29925 (BRI); D,F from Bean 30027
(BRI); E from Hines s.n. (BRI [AQ779478]).

Solanum cleistogamum Symon, Trans.
& Proc. Roy. Soc. South Australia 95: 227
(1971). Type: Western Australia, c. 32 km
N of Onslow, 1 July 1967, D.E. Symon 5418
(holo: PERTH; iso: AD, CANB, K, L).

S. petrophilum var. pedicellatum Ewart &
Davies, FI. Northern Territory 243 (1917).
Type: Northern Territory. Lat. 19° long.
132°, 4 July 1911, G.F. Hill 435 (holo: MEL
[2 sheets]).

420

Illustrations : Cunningham et al. (1981: 592);
Symon(1981: 194)

Herbaceous resprouter, 0.15-0.25 m high.
Stems initially ascending, then prostrate
and trailing. Adult branchlets white to
rusty-brown; prickles 7-18 per cm, straight,
acicular, 1-6 mm long, 15-20 times longer
than wide, glabrous; stellate hairs dense
or very dense, lateral rays 8-15, porrect,
ascending or multiradiate, central ray 1-
2.5 times as long as laterals. Adult leaves
lanceolate or ovate, entire or with 2 or 3
shallow obtuse lobes on each side, lobing
index 1-1.1; lamina 2.8-47 cm long, 1.2-3.6
cm wide, 1.2-2.4 times longer than broad,
apex obtuse, base obtuse or cordate, oblique
part 0-4.5 mm long, obliqueness index
0-10%; petioles 1.4-3.1 cm long, 40-70%
length of lamina, prickles present. Upper leaf
surface grey-green, prickles absent; stellate
hairs moderately dense or dense, stalks 0-0.1
mm long, lateral rays 6-8, porrect, central
ray 1.1-2 times as long as laterals. Lower
leaf surface green or greenish-white, prickles
absent or with 1-10 on midvein only, straight,
acicular. Lower leaf surface stellate hairs
moderately dense or dense, stalks 0-0.1 mm
long, lateral rays 7-8, porrect; central ray 1.2-
2 times as long as laterals, sometimes gland-
tipped. Inflorescence supra-axillary, cymose
(pseudo-racemose), common peduncle

8- 21 mm long, rachis prickles present, 1-5-
flowered, with all flowers bisexual. Flowers
5-merous. Pedicels at anthesis 7-14 mm long,
prickles present. Calyx tube at anthesis 2-3.5
mm long; lobes deflate or rostrate, 2-4 mm
long; prickles present, 10-25 per flower,
1.5-3 mm long; stellate hairs dense, lateral
rays 6-8; central ray 1.2-2.2 times as long
as laterals, sometimes gland-tipped. Corolla
rotate, 6-9 mm long, white or mauve, inner
surface glabrous; anthers 2-2.8 mm long;
filaments 1.2-1.3 mm long; ovary glabrous;
style 5-6 mm long, erect near base, then
bent, displacing one anther, glabrous; stigma
slightly expanded, obscurely lobed. Fruiting
calyx lobes less than or more than half length
of mature fruit, prickles 1.5-5 mm long.
Mature fruits 1-3 per inflorescence, globular,

9- 12 mm diameter, yellow, 2-locular; placenta
sessile, semi-circular; interior juicy, pericarp

Austrobaileya 8(3): 412-430 (2011)

0.4-0.8 mm thick; pedicels 15-23 mm long.
Seeds 2-2.5 mm long, white or pale yellow.

Additional selected specimens examined: Western
Australia. Horridum Hill, near highway, c. 50 km E
of Port Hedland, Apr 2006, Bean 25014 & Lepschi
(BRI, CANB, PERTH). Northern Territory. Track
to Lake Lewis, S of Tilmouth Bore, c. 160 km NW
of Alice Springs, Aug 2006, Bean 25472 & Albrecht
(BM, BRI, CANB, MEL, NSW, NT, NY); North-west
Simpson Desert, Sep 1973, Latz 4658 (AD, BRI, NT);
Curlew Waterhole, Lander River, 70 miles [113 km] NW
of Willowra Homestead, Central Australia, Jul 1958,
Chippendale 4799 (BRI, NT); W margin of Lake Ruth, c.
5 km SE of Mongrel Downs Homestead, Tanami Desert,
Aug 1978, Donner 6283 (AD, BRI). Queensland. Burke
District: Roxmere Station, S of Cloncurry, Apr 2003,
Booth 3168 & Kelman (AD, BRI, NSW). South Kennedy
District: Lake Buchanan, May 1980, Hyland 10525
(BRI, CNS). Warrego District: Thargomindah, Jun
1936, Blake 11781 (BRI); 10.9 km from Adavale towards
Blackall at tank, Oct 1983, Canning 6206 (BRI, CANB).
Maranoa District: Karoola Road, S of Dirranbandi,
10 km N of state border, Sep 2004, Bean 22847 (BRI).
New South Wales. 3.6 km NE of Cumborah, towards
Lightning Ridge, Sep 2004, Bean 22893 (BRI); Fowlers
Gap, N of Broken Hill, Oct 1975, Jacobs 2337 (BRI,
NSW); ‘Tundulya’, c. 25 miles [40 km] SE of Louth, Apr
1967, Moore 4949 (CANB). South Australia. E side of
Lincoln Highway, 17 km N of Whyalla, Oct 1998, Jobson
5736 & Downs (BRI, NSW).

Distribution and habitat: Solarium
cleistogamum is widespread in mainland
Australia south of about latitude 19°, from the
west coast of Western Australia to the slopes
of the Great Divide in New South Wales and
Queensland. In South Australia, it occurs as
far south as Spencer Gulf.

It grows on a range of habitats but prefers
sandy or sandy-loam soils, often on rather
low-lying terrain, but also sometimes on steep
hillsides.

Phenology: Flowers and fruits may be found
throughout the year.

Notes: This species was previously included
in the synonymy of Solarium ellipticum (Bean
2004). I now recognise that this was incorrect,
and that the two are distinct. While the two
species are closely related and rather difficult
to distinguish in the herbarium, the relatively
small tasty yellow fruits, the small corolla, and
the ± cordate leaf bases of S. cleistogamum
readily distinguish it.

Bean, Solarium ellipticum group

Solanum ellipticum R.Br., Prodr. 446
(1810); S. ellipticum var. ellipticum , Benth.,
FI. Austral. 4: 464 (1868); S. ellipticum f.
ellipticum , Wawra, Itin. Princ. S. Coburgi
100 (1883); S. ellipticum var. typicum Domin,
Biblioth. Bot. 89: 588 (1929), nom. inval.
Type: [Queensland. Port Curtis District:]
‘Broadsound’, 25 September 1802, R.
Brown [Bennett Number 2674] (lecto: BM
[000596888]; isolecto: MPU, fide Symon
[1981: 188]).

Herbaceous resprouter, or rhizomatous
perennial shrub 0.1-0.25 m high. Stems
initially ascending, then prostrate and
trailing. Adult branchlets grey, purple, yellow
or brown; prickles 1-10 per cm, straight,
acicular, 1-8 mm long, 8-14 times longer
than wide, glabrous or with dense stellate
hairs on lower part; stellate hairs dense to
very dense, 0.5-0.8 mm diameter, stalks
0-0.1 mm long, lateral rays 7-10, lateral rays
porrect or ascending, central ray 0.5-1.2 times
as long as laterals. Adult leaves elliptical or
ovate, entire; lamina 3.5-14 cm long, 1.5-5.2
cm wide, 1.7-2.9 times longer than broad,
apex obtuse or acute, base cuneate or obtuse,
oblique part 0-9 mm long, obliqueness index
0-6%; petioles 0.8-4 cm long, 20-45% length
of lamina, prickles absent or present. Upper
leaf surface green or grey-green, prickles
2-20, present on midvein only or on midvein
and lateral veins, straight, acicular, 1-7 mm
long; stellate hairs distributed throughout,
sparse to dense, 0.1-0.5 mm apart, 0.4-0.7
mm across, stalks 0-0.1 mm long, lateral rays
7-8, porrect, central ray 0.5-1 times as long
as laterals; simple hairs absent; Type 2 hairs
absent. Lower leaf surface greenish-white to
grey or yellowish; prickles 0-20, absent or
present on midvein only or on midvein and
lateral veins, straight, acicular; stellate hairs
moderately dense to very dense, 0.05-0.3
mm apart, 0.5-0.8 mm diameter, stalks 0-0.2
mm long, lateral rays 7-8, porrect, central ray
0.5-1.2 times as long as laterals; simple hairs
absent; Type 2 hairs absent. Inflorescence
supra-axillary, solitary or cymose (pseudo-
racemose), common peduncle 0-6 mm long,
rachis prickles present, 1-5-flowered, with
some bisexual and some male flowers. Flowers
5-merous. Pedicels at anthesis 4-11 mm long,

421

0.7-0.9 mm thick at mid-point, prickles
absent or present. Calyx tube at anthesis 1.5-

3 mm long; lobes deflate or attenuate, 2-6
mm long; prickles 5-50 per flower, 1-5 mm
long; stellate hairs moderately dense to very
dense, white to yellow or rusty, 0.4-0.8 mm
across, stalks 0-0.3 mm long, lateral rays 7-
8, central ray 0.8-1.3 times as long as laterals;
simple hairs absent; Type 2 hairs absent.
Corolla shallowly or deeply lobed, 7-12 mm
long, mauve or purple, inner surface with a
few stellate hairs near lobe apices. Anthers
4-5.5 mm long. Ovary glabrous or with Type
2 hairs only; functional style 5.5-10 mm long,
protruding between anthers, glabrous or with
Type 2 hairs only; stigma unexpanded, same
colour as style. Fruiting calyx lobes less than
or more than half length of mature fruit,
prickles 1-5 mm long. Mature fruits 1-2 per
inflorescence, globular, 13-17 mm diameter,
yellowish-green, 2-locular; placenta stalked,
anvil-shaped; interior moist but not juicy,
pericarp 0.6-1 mm thick; pedicels 7—19 mm
long, 0.8-2.1 mm thick at mid-point. Seeds
2.2-27 mm long, pale yellow. Fig. 1.

Additional selected specimens examined : Queensland.

Leichhardt District: Slopes of Marling Spike, Homevale
N.P., NNW of Nebo, Dec 2010, Bean 30650 et al. (BRI).
Port Curtis District: Orange Creek, c. 20 miles [32
km] NW of Biloela, Jun 1959, Johnson 857 (BRI); 4
km WSW of St Lawrence, Apr 2000, Bean 16250 (BRI,
MEL). Burnett District: ‘Neaavie’, NE of homestead
towards Barambah Creek, Aug 1996, Grimshaw PG2536
& Turpin (BRI); c. 6 km NE of Allies Creek and 75 km
SW of ‘Weir Weir’, Jul 1998, Pollock ABP617 & Dean
(BRI). Wide Bay District: Bruce Highway, between
Gin Gin and Miriam Vale, Jan 2006, Bohs 3540 & Bean
(BRI); Bingera, Dec 1896, Bailey s.n. (BRI [AQ39062]).
Darling Downs District: 8 km WSW of Oakey, Apr
1994, Fensham 1444 (BRI); 14 km N of Goondiwindi,
towards Moonie, Feb 1996, Bean 9907 (BRI, MEL,
NSW). Moreton District: Gatton, Nov 1916, Bick s.n.
(BRI [AQ39059]); 2 km ESE of Laidley, Jun 2002, Bean
19062 (BRI). New South Wales. North West Plains:

4 km SSW of ‘The Glen’ Homestead, c. 70 km W of
Moree, Jan 1999, Wannan 1036 et al (AD, BRI, NSW).

Distribution and habitat : The limits of
distribution of Solanum ellipticum sens. str.
are not yet known with certainty, but it is
found in the Queensland Pastoral districts of
Moreton, Darling Downs, Wide Bay, Burnett,
Leichhardt and Port Curtis. It also occurs on
the north-western plains of New South Wales
near Moree.

422

Phenology : Flowers and fruits can be found
throughout the year.

Notes : Solanum ellipticum is a prostrate to
semi-erect plant with trailing stems (Fig. 1).
It has a small inflorescence with 1-5 flowers,
and some of these are functionally male
(styles very short and non-functional). It is
perhaps the only species of the group in which
the flowers are not all bisexual. The common
peduncle is usually absent, although it can be
up to 6 mm long. The number of fruits is one
or two.

Solanum ellipticum var. chillagoense
Domin, S. ellipticum f. albiflora Domin, and
S. sp. (Newcastle Range D.E. Symon 4907)
were included by Bean (2004) in the synonymy
of S. ellipticum. The author now recognises
that this is incorrect; however, further study
is required to accurately place these names
beyond synonymy with S. ellipticum.

Solanum emmottii A.R.Bean species nova

S. lithophilo affinis sed ab eo corolla minore,
calycis aculeis paucioribus brevioribusque
et pilis stellatis omnibus stipitibus quam 0.2
millimetris longitudine brevioribus praeditis
differens. Typus: Queensland. Gregory
North District: Near homestead, Ethabuka
Reserve, 8 November 2010, A. J. Emmott B690
(holo: BRI [1 sheet + spirit]; iso: AD, NT, NY,
W, distribuendi ).

Rhizomatous perennial shrub 0.2-0.5 m
high. Stems initially sprawling, then more
or less erect. Adult branchlets white, grey
or yellowish; prickles 1-28 per cm, straight,
acicular, 1-5 mm long, 10-14 times longer
than wide, glabrous; stellate hairs very dense,
0.4-0.8 mm diameter, stalks 0-0.2 mm
long, lateral rays 7-13, porrect, ascending
or multiradiate, central ray 0.8-1.3 times as
long as laterals; Type 2 hairs absent. Adult
leaves ovate, entire; lamina 3.5-8.8 cm long,
1.4-3.8 cm wide, 1.7-2.9 times longer than
broad, apex obtuse or acute, base cuneate,
obtuse or cordate, oblique part 0-4 mm long,
obliqueness index 0-7%; petioles 0.6-4.2
cm long, 18-48% length of lamina, prickles
absent or present. Upper leaf surface grey-
green or grey, prickles absent; stellate hairs
dense to very dense, 0.05-0.2 mm apart, 0.4-
0.7 mm across, stalks 0-0.1 mm long, lateral

Austrobaileya 8(3): 412-430 (2011)

rays 6-10, porrect, central ray 0.7-1.2 times
as long as laterals; simple hairs absent; Type 2
hairs absent. Lower leaf surface white or grey;
prickles absent or present on midvein only,
0-2, straight, acicular; stellate hairs dense to
very dense, 0.05-0.1 mm apart, 0.4-0.6 mm
diameter, stalks 0-0.3 mm long, lateral rays
7-11, porrect or ascending, central ray 0.7-1.2
times as long as laterals; simple hairs absent;
Type 2 hairs absent. Inflorescence supra-
axillary, cymose (pseudo-racemose), common
peduncle 15-50 mm long, rachis prickles
absent or present, 2-6-flowered, with all
flowers bisexual. Flowers 5-merous; pedicels
at anthesis 4-15 mm long, same thickness
throughout, prickles absent or present; calyx
tube at anthesis 2.5-4 mm long; lobes deflate
or attenuate, 2-7 mm long; prickles 1-25 per
flower, 0.5-2 mm long; stellate hairs very
dense, yellow or white, 0.4-0.8 mm across,
stalks 0-0.3 mm long, lateral rays 8-10,
central ray 0.7-1.3 times as long as laterals;
simple hairs absent; Type 2 hairs absent.
Corolla rotate or shallowly lobed, 9-13 mm
long, purple, inner surface sparsely stellate-
hairy; anthers 4.2-5.5 mm long; filaments
1.2-1.6 mm long. Ovary glabrous; functional
style 8-9.5 mm long, strongly bent near base,
not touching anthers, eccentric, glabrous or
rarely with stellate hairs; stigma expanded,
darker than style, obscurely lobed. Fruiting
calyx lobes less than or more than half
length of mature fruit. Mature fruits 1-3 per
inflorescence, globular, c. 13 mm diameter,
colour unknown, 2-locular; placenta sessile,
semicircular; interior moist but not juicy,
pericarp 0.3-0.4 mm thick; pedicels 8-13 mm
long, 1.2-1.5 mm thick at mid-point. Seeds
2.5-3.3 mm long, pale yellow. Fig. 5.

Additional specimens examined : Northern Territory.
c. 5 km east of Lake Caroline, northern Simpson
Desert, Jul 2007, Albrecht 12300 & Duguid (BRI,
NT); c. 1 km west of Lake Caroline, northern Simpson
Desert, Jul 2007, Albrecht 12340 & Duguid (BRI, NT);
Near Hale River, Simpson Desert, Sep 1955, Perry
5421 (BRI, CANB); 12 miles [20 km] NW of Lucy
Creek Homestead, Jul 1957, Chippendale 3523 (BRI).
Queensland. Mitchell District: Valetta Bore paddock,
‘Valetta’, near ‘Noonbah’, c. 170 km SW of Longreach,
May 2010, Bean 29753 & Emmott (AD, BM, BRI, NSW,
NT); south boundary of Valetta Bore paddock, ‘Valetta’,
c. 180km SW of Longreach, May 2010, Bean 29757 &
Emmott (BRI, MEL). Gregory North District: 10 km
NNW of Bedourie, (1.6 km W of Duck Hole), Kamaran
Downs Station, Jun 1995, Edmunds 23 (BRI). Gregory

Bean, Solanum ellipticum group

423

Fig. 5. Solanum emmottii. A. flowering branchlet x 1.5. B. lateral view of flower x 4. c. stamens, style and ovary x 4.
D. immature fruit x 2. All from Bean 29753 & Emmott (BRI).

South District: Mt Howitt Station, 80 miles [133 km] W
of Eromanga, Jul 1936, Blake 11926 (BRI); 10 miles [16
km] W of Betoota, Jul 1936, Blake 12184 (BRI); ‘Kyra’,
40 miles [67 km] W of Quilpie, Jul 1965, Emmerson s.n.
(BRI [AQ39068]); 35 km W of Betoota on Birdsville
road, Aug 1979, Williams 78187 (BRI). South Australia.
54 km by road S of Birdsville on the Birdsville Track to
Marree, Mar 2001, Thomas 2049 & Fechner (BRI).

Distribution andhabitat: Solanum emmottii is
known from the far south-west of Queensland,
and adjacent areas of Northern Territory and
South Australia (Map 3). It grows almost
exclusively on or adjacent to sand dunes,
in red to orange sandy soil, associated with
shrubs such as Crotalaria eremaea F.Muell.
However, at the most north-easterly location,

it occurs on the slopes of gravelly hills in
brown or yellow loam, and in association with
Atalaya hemiglauca (F.Muell.) F.Muell. ex
Benth., Acacia ensifolia Pedley, Eremophila
mitchellii Benth. and Triodia longiceps
J.M.Black.

Phenology : Flowers have been recorded from
March to November; a potentially mature
fruit was collected in July.

Notes : Solanum emmottii differs from
S. lithophilum by the non-floccose branchlet
indumentum and the stellate hairs stalks less
than 0.2 mm (floccose branchlet indumentum
with stalks 0-2 mm long for S. lithophilum );

424

stellate hairs with shorter central rays; shorter
calyx lobes; calyx prickles 1-25, each 0.5-2
mm long (25-50 calyx prickles, each 2-4 mm
long for S. lithophilum ); corolla 9-13 mm long
(16-20 mm for S. lithophilum ); the presence
of stellate hairs on the inner corolla surface
(glabrous for S. lithophilum ); and the smaller
fruits.

Edmunds 23 (cited above) differs from
the other cited specimens by the presence of
numerous stellate hairs on the style, but is in
other regards quite typical.

Conservation status : This species appears
to be fairly common and is not currently
considered to be under threat. A conservation
status of ‘Least Concern’ is recommended.

Etymology : The species is named for Angus
James Emmott (1962-), grazier, accomplished
naturalist, and dedicated advocate for the
preservation of natural ecosystems in arid
Australia.

Solanum lithophilum F.Muell., Linnaea
25: 434 (1853). Type: South Australia. Near
Cudnaka [Kanyaka Homestead], Flinders
Ranges, October 1851, F. Mueller s.n. (lecto:
MEF11873 ,fide Symon [1981: 188]).

S. ellipticum var. mollibaccalis J.M.Black,
Trans. & Proc. Roy. Soc. South Australia 52:
227 (1928). Type: South Australia. Finniss
Springs near Fake Eyre, s.dat., F.D.Warren
s.n. (syn: AD, fide Symon [1981: 188], n.v).

S. ellipticum var. Foot hills (G.J.Leach 1145)-,
Kerrigan & Albrecht (2007), Albrecht et al.
(2007: 176).

S. ellipticum R.Br. auct. non Symon (1981),
pro parte.

Herbaceous resprouter, 0.3-0.45 m high.
Stems initially sprawling, then more or less
erect. Adult branchlets grey, yellow or rusty;
prickles 9-40 per cm, straight, acicular,
2-7 mm long, 13-18 times longer than wide,
glabrous; stellate hairs very dense, stalks
0-2 mm long, lateral rays 8—13, porrect or
multiradiate, central ray 1.1-2.2 times as long
as laterals. Adult leaves elliptical or ovate,
entire; lamina 3.6-10.1 cm long, 1.5-5.3 cm
wide, 1.9-3.8 times longer than broad, apex
acute, base cuneate or obtuse, oblique part 0-

Austrobaileya 8(3): 412-430 (2011)

6 mm long, obliqueness index 0-7%; petioles
1.3-6 cm long, 33-60% length of lamina,
prickles present. Upper leaf surface grey-
green or grey; prickles absent or present on
midvein only, 0-2, straight, acicular, 3-7 mm
long; stellate hairs dense, stalks 0-2 mm long,
lateral rays 7-10, porrect or ascending, central
ray 1.2-2 times as long as laterals. Fower leaf
surface white, grey or silvery; prickles absent
or present on midvein only, 0-4, straight,
acicular; stellate hairs very dense, lateral
rays 8-15, porrect or multiradiate, central ray
1.2-2 times as long as laterals. Inflorescence
supra-axillary, cymose (pseudo-racemose),
common peduncle 20-52 mm long, rachis
prickles present, 3-7-flowered, with all
flowers bisexual. Flowers 5-merous. Pedicels
at anthesis 7-16 mm long, same thickness
throughout, prickles absent or present. Calyx
tube at anthesis 3-4 mm long; lobes attenuate,
5-11 mm long; prickles present, 25-50 per
flower, 2-4 mm long; stellate hairs very
dense, yellowish-white or purple, lateral rays
8-13, central ray 1-2 times as long as laterals.
Corolla rotate, 14-20 mm long, purple, inner
surface glabrous. Anthers 4.7-6 mm long.
Filaments 1-1.2 mm long. Ovary glabrous;
style 9-13 mm long, strongly bent near base,
not touching anthers, eccentric, glabrous;
stigma expanded, darker than style, obscurely
lobed. Fruiting calyx lobes less than or more
than half length of mature fruit, prickles 2.5-4
mm long. Mature fruits 1-2 per inflorescence,
globular or ellipsoidal, 18-21 mm diameter,
purplish-green or almost white, 2-locular;
placenta stalked, anvil-shaped; interior
juicy, succulent; pericarp 0.3-0.8 mm thick;
pedicels 12-25 mm long. Seeds 2.8-3.2 mm
long, pale yellow or white. Figs. 2 & 6.

Additional selected specimens examined: Western
Australia. 100 miles [161 km] N ofEucla, s.dat., Batts.n.
(MEL). Northern Territory. Stuart Highway, 10.7 km N
of Aileron, Aug 2006, Bean 25457 & Albrecht (AD, BRI,
NT); track to Lake Lewis, S of Tilmouth Bore, c. 160 km
NW of Alice Springs, Aug 2006, Bean 25473 & Albrecht
(BRI, NT); Western side of Kings Canyon Range, 1 km
south of Carmichael Crag, Aug 1986, Menkhorst s.n.
(MEL); Orange Creek, 1 km S of Homestead on Stuart
Highway, 24 km S of Alice Springs on Adelaide Road,
Jun 1968, Must 17 (MEL); 4 miles [6 km] W of Central Mt
Wedge Homestead, Jun 1955, Chippendale 1283 (BRI);
Uluru N.P.: Kata Tjuta, on the Docker River Road, 35.7
km WNW of Ranger Station, May 1988, Lazarides &
Palmer 108 (CANB); Beside Todd River, N of Schwarz

Bean, Solatium ellipticum group

Crescent, Alice Springs, Aug 2006, Bean 25494 (BRI,
CANB, NT); N’Dahla Gorge, Ross River area, E
MacDonnell Ranges, Jun 1974, Williss.n. (MEL); S bank
of upper Finke River opposite Mt Sonder, Jul 1966, Willis
& Morrison s.n. (MEL). Queensland. Gregory South
District: on track 13.4 km due NW of ‘Old Karmona’
cottage, Aug 2008, Turpin GPT1280 & May (BRI, CNS);
on track 26.9 km due NW of Orientos Station, Aug 2008,
Turpin GPT1279 & May (BRI, CNS). New South Wales.
Tarella, Aug 1887, Baeuerlen 118 (MEL); Golden Gully
mining area adjacent to the Deadhorse Gully camping
area, 1 km N of Tibooburra in Sturt N.R, Sep 1989,
Coveny 13598 et al. (AD, BRI, MO, NSW); ridge at end
of Mootwingee Gorge Road, Mootwingee N.R, Aug
1989, Porteners 33 & Plaza (NSW, NT); 2 km W of Big
Wallaby tank, 5 km NNW of Koonenberry Mountain,
Sep 1971, De Nardi 847 (NSW); Silverton near Barrier
Range, Nov 1884, Harris s.n. (MEL); Mt Koonenberry,
Dec 1860 , Beckler s.n. (MEL). South Australia. nearMt
Everard, 1882, Giles s.n. (MEL); Fogarty’s claypan. Mar
1984, Jackson 5255 (MEL); Strangways Springs, 1895,
Gratwick 16 (MEL); NW of Ooldea, s.dat., Tietkins s.n.
(MEL); Brachina Creek, Jun 1960, Filson 2115 (MEL); 2
miles [3.3 km] S of Emu, Aug 1956, Forde 389 (MEL);
Innamincka Homestead, Jun 1980, Lay 1493 (MEL);
Lake Torrens Basin, east Mernmerna, c. 150 km NE of
Port Augusta, on road to Marree, Oct 1968, Weber 729
(MEL).

Distribution and habitat : Solanum
lithophilum is widespread in South
Australia (except in the south-east) and
the southern one-third of the Northern
Territory. It also extends to far eastern
parts of Western Australia and north¬
western New South Wales and it has
recently been recorded from the extreme
south-west of Queensland (Map 3). In
South Australia, it is frequently recorded
from rocky hill slopes, but in the Northern
Territory it often grows on rather sandy
colluvial and alluvial soils, with only
limited occurrences on steep hillsides.

Phenology : Flowers are recorded from all
months of the year; fruits are recorded from
March to September.

Affinities: Solanum lithophilum may be readily
distinguished from S. ellipticum. Solanum
lithophilum is a multi-stemmed plant to 45
cm high; the stems are at first sprawling, but
finally erect and have a floccose tomentum;
the lower leaves have very long petioles, the
inflorescences are well displayed and the very
showy flowers (corolla 14-20 mm long) have
a prominently eccentric style. The fresh fruits
are large, 18-21 mm in diameter, and often

425

bone-white when mature, at least at the distal
end.

By contrast, Solanum ellipticum rarely
exceeds 20 cm in height, the stems are
ascending near the base, but then prostrate
or procumbent, the tomentum is not floccose,
all leaves have petioles of a similar length,
the inflorescences are not well displayed and
the flowers could not be described as showy
(corolla 7-12 mm long). The style is erect
rather than eccentric, protruding between
the anthers. The fruits are smaller, 13-17
mm across, and green to yellow-green at
maturity.

There are further differences at the
microscopic level. Some stellate hairs on
the branchlets of Solanum lithophilum are
multiradiate with 10-13 lateral rays, and
the central ray on all plant parts is 1.1-2.2
times longer than the laterals. The stigma
of Solanum lithophilum is conspicuously
swollen, and green in colour (cf. S. ellipticum
where the stigma is scarcely broader than the
style, and is the same colour as it).

Notes: Well-developed plants of Solanum
lithophilum form a small multi-stemmed
bush with numerous, large, well-displayed
purple flowers (Fig. 2). Such plants are very
attractive and have considerable horticultural
potential.

Solanum unispinum A.R.Bean species
nova S. cleistogamo affinis sed ab eo corolla
profunde lobata, antheribus longioribus, radio
centrali pilorum stellatorum breviore, calyce
aculeis pauculis vestito et habitu fruticoso
differens. Typus: Queensland. Gregory
North District: Winton - Jundah Road, 2.7
km N of Mayne River crossing, 25 May 2004,
A.R. Bean 22512 (holo: BRI; iso: CANB,
NSW, distribuendi).

Herbaceous resprouter or rhizomatous
perennial shrub, to 0.3 m high. Stems initially
sprawling, then more or less erect. Adult
branchlets purplish, yellow or rusty. Adult
branchlet prickles 1-4 per cm, straight,
acicular, 3-8 mm long, 12-16 times longer
than wide, glabrous; stellate hairs very dense,
0.4-0.7 mm diameter, stalks 0-0.1 mm
long; lateral rays 8-10, porrect; central ray

426

Austrobaileya 8(3): 412-430 (2011)

Fig. 6. Solatium lithophilum. A. flowering branchlet x 0.8. B. leaf from lower part of plant * 0.8. C. oblique view of an
open flower x 2. D. lateral view of a flower bud x 2. E. mature fruit with attached calyx x 1.5. F. cross-section of mature
fruit x 2. A, E, F from Bean 25457 & Albrecht (BR1); B-D from Bean 25494 (BRI).

427

Bean, Solanum ellipticum group

0.4-0.8 times as long as laterals, not gland-
tipped. Adult leaves ovate or broadly ovate,
entire; lamina 2.2-3.5 cm long, 1.4-2.1 cm
wide, 1.6-1.7 times longer than broad, apex
obtuse or acute, base obtuse, oblique part
0-2 mm long, obliqueness index 0-9%.
Petioles 1.4-2.7 cm long, 58-77% length of
lamina, prickles present. Upper leaf surface
grey-green, prickles absent; stellate hairs
distributed throughout, dense, 0.15-0.2 mm
apart, 0.4-0.5 mm across, stalks 0-0.1 mm
long, lateral rays 8, central ray 0.6-0.9 times
as long as laterals, not gland-tipped; simple
hairs absent; Type 2 hairs absent. Lower
leaf surface white or grey; prickles absent
or present on midvein only, 0-3, straight,
acicular; stellate hairs dense or very dense,
0.05-0.1 mm apart, 0.4-0.55 mm diameter,
stalks 0-0.1 mm long, lateral rays 8-9, porrect;
central ray 0.4-1 times as long as laterals, not
gland-tipped; simple hairs absent; Type 2 hairs
absent. Inflorescence supra-axillary, cymose
(pseudo-racemose), common peduncle 3-10
mm long, rachis prickles absent or present, 1-
2-flowered, with all flowers bisexual. Flowers
5-merous. Pedicels at anthesis 6-13 mm long,
same thickness throughout, 0.5-0.6 mm thick
at mid-point, prickles absent. Calyx tube at
anthesis 2.5-3 mm long; lobes attenuate, 1.5-
2 mm long. Calyx prickles absent or present,
0-4 per flower, 1-4 mm long. Calyx stellate
hairs very dense, yellow or white or purple,
0.3-0.5 mm across, stalks absent, lateral rays
7-8, central ray 0.5-1 times as long as laterals,
not gland-tipped; simple hairs absent; Type 2
hairs absent. Corolla deeply lobed, 7-10 mm
long, purple, inner surface glabrous. Anthers
37-4.6 mm long. Ovary glabrous; style 7.5-
9 mm long, straight, protruding between
anthers or slightly bent, displacing one anther,
glabrous; stigma expanded, yellow, obscurely
lobed. Fruiting calyx lobes less than or more
than half length of mature fruit; prickles
absent or present, 2-4 mm long. Mature
fruits 1-2 per inflorescence, globular, 11-13
mm diameter, yellowish-green, 2-locular;
placenta sessile elliptical to semi-circular;
interior juicy, succulent; pericarp 0.2-0.5 mm
thick; pedicels 12-20 mm long. Seeds 2.5-27
mm long, pale yellow. Fig. 7.

Additional specimens examined: Queensland.

Mitchell District: c. 160 km S of Longreach, Apr 1989,

Emmott 286 (BRI); Noonbah Station, in Back Paddock,
SW corner of Back Paddock, Jan 2008, Emmott B516
(BRI, CANB); SW corner of Back Paddock, ‘Noonbah’,
c. 170 km SW of Longreach, May 2010, Bean 29771 &
Emmott (BRI); SW corner of Back Paddock, ‘Noonbah’,
c. 170 km SW of Longreach, May 2010, Bean 29772 &
Emmott (BRI, MEL). Gregory North District: 86.6 km
W of Winton on the Winton - Boulia Road, Nov 2010,
Emmott B682 & Denny { BRI, NY); 98.9 km W of Winton
on the Winton - Boulia Road, Nov 2010, Emmott B683 &
Denny (BM, BRI); Winton - Jundah Road, 4.4 km S of
‘Elvo’ Homestead, May 2004, Bean 22553 (BRI).

Distribution and habitat : Endemic to south¬
western Queensland between Stonehenge
and Winton (Map 1). It grows in shrubland
or low open woodland, on stony hills and
ranges, with shallow sandy soil. Associated
species include Eucalyptus normantonensis
Maiden & Cambage, Acacia shirleyi Maiden,
A. aneura and Triodia sp.

Phenology: Flowers have been recorded in
April, May and November; fruits in January,
May and November.

Notes: Solanum unispinum is similar in
appearance to S. cleistogamum , but differs
by the bushy habit and ascending branches,
branchlet prickles 1-4/cm (7-18/cm for S.
cleistogamum ), the central ray of the stellate
hairs 0.4-1 times as long as laterals (1-2.5
times for S. cleistogamum ), the leaf bases not
cordate, the calyx prickles 0-5 (10-25 for S.
cleistogamum ), the deeply lobed corolla (rotate
for S. cleistogamum ), and the anthers 37-4.6
mm long (2-2.8 mm for S. cleistogamum).

Conservation status: At the two sites on the
Winton - Boulia road, four plants and one
plant were found respectively. At ‘Noonbah’,
it is known from fewer than 20 plants. Only
two plants were seen near ‘Elvo’ Homestead,
while at the type locality there were fewer
than 10 plants. Using the criteria of the IUCN
Red List (IUCN 2000), a conservation status
of Critically Endangered (Criterion D) is
recommended.

Etymology: From the Latin uni meaning one
and spinus meaning spine. The calyces of this
species have no more than a single prickle on
each lobe, and sometimes only one calyx lobe
bears a prickle.

428

Austrobaileya 8(3): 412-430 (2011)

Fig. 7. Solanum unispinum. A. flowering branchlet x 1. B. oblique view of an open flower x 3. C. lateral view of flower
x 3. C. stamens, style and ovary x 4. D. cross-section of mature fruit x 3. All from Bean 22512 (BRI).

Acknowledgements

I am grateful to David Albrecht (NT) for taking
me into the field around Alice Springs to study
various Solanum taxa, and for convincing me
that S. cleistogamum is a distinct species.
Angus Emmott of Noonbah Station kindly
escorted me around his property to see
noteworthy solanums that he had found, and
supplied many images of them; he has also
made excellent collections from other parts

of western Queensland at my request. Will
Smith (BRI) provided the illustrations and
the distribution maps. Peter Bostock expertly
translated the diagnoses into Latin.

Bean, Solarium ellipticum group

References

429

Albrecht, D.E., Duguid, A.W., Coulson, H., Harris,
M.G. & Latz, P.K. (2007). Vascular Plant
Checklist for the Southern Bioregions of the
Northern Territory , 2nd edition. Northern
Territory Government: Alice Springs.

Bean, A.R. (2004). The taxonomy and ecology of
Solanum subg. Leptostemonum (Dunal) Bitter
(Solanaceae) in Queensland and far north¬
eastern New South Wales. Austrobaileya 6:
639-816.

Brooke, G. & Mcgarva, L. (1998). The Glove Box
Guide to plants of the NSW rangelands. NSW
Agriculture: Sydney.

Cunningham, G.M., Mulham, W.E., Milthorpe, P.L. &
Leigh, J.H. (1981). Plants of Western New South
Wales. Soil Conservation Service and N.S.W.
Government Printing Office: Sydney.

Henderson, R.J.F. (2002). Names and Distribution
of Queensland Plants, Algae and Lichens.
Queensland Herbarium, Environmental
Protection Agency: Toowong.

Iucn (2001). Red List Categories and Criteria: Version
3.1. IUCN Species Survival commission. IUCN:
Gland, Switzerland/Cambridge, U.K.

Kerrigan, R.A. & Albrecht, D.E. (2007). Checklist
of NT Vascular species, http://www.nt.gov.au/
nreta/wildlife/plants/pdf/200701nt_checklist.

pdf Accessed 17 th October 2010.

Symon, D.E. (1981). A revision of the genus Solanum
in Australia. Journal of the Adelaide Botanic
Gardens 4: 1-367.

Map 1 . Distribution of Solanum aridicola • and S. unispinum ▲

430

Austrobaileya 8(3): 412-430 (2011)

Map 2. Distribution of Solarium callosum

Map 3. Distribution of Solarium emmottii ▲ and S. lithophilum •

Austrobaileya 8(3): 431-434 (2011)

431

SHORT COMMUNICATION

Luvunga monophylla (DC.) Mabb. (Rutaceae):
a new species for Queensland

David G. Fell 13 & David J. Stanton 2

'3D Environmental, P.O. Box 337, Alstonville NSW 2477, Australia. Email:
davidfell@3denvironmental.com.au; Corresponding author

2 3D Environmental, PO Box 959, Kenmore, Queensland 4069, Australia.

The genus Luvunga Buch.-Ham. ex Wight
& Arn. contains twelve species occurring
in Asia, Malesia and Australia (Hyland et
al. 2003). Luvunga monophylla is the sole
Australian representative of Rutaceae subtribe
Triphasiineae in the tribe Aurantieae Rchb.
(Mabberley 1998); however, infrafamilial
relationships in this part of the family remain
to be fully resolved utilising molecular data
and this group may not be recognized in the
future (Kubitzki et al. 2010).

Luvunga monophylla (DC.) Mabb. is
a facultatively deciduous, spiny shrub or
climber, 0.5-3 m high with white, cream,
yellow flowers, obovoid fleshy fruit to 10 mm
diameter, and one or two seeds per fruit (Pedley
1987; Russell-Smith & Dunlop 1992; Hyland
et al. 2003; Department of Environment and
Conservation 2010). It was first collected from
Australia in the Sir Edward Pellew Group of
islands, Gulf of Carpentaria, in December
1802 by Robert Brown during Flinders’
Investigator circumnavigation of Australia
and was collected in Timor by J.B.L.C.T.
Leschenault de la Tour during Baudin’s
1800-1803 expedition in Le Geographe and
Le Naturaliste (Mabberley 1998).

It has now been recorded on lama (Yam
Island), Torres Strait which is the first record
of the species for Queensland. Prior to this
collection it had been recorded from Australia
in the North Kimberley and Dampierland
bioregions of Western Australia (Department
of Environment & Conservation 2010) and in
the Arnhem Coast, Central Arnhem, Tiwi,
Gulf Coastal, Gulf Fall and Uplands and

Darwin Coastal Bioregions of the Northern
Territory. The habitat in these areas includes
monsoon forest, deciduous vine thicket and
semi-deciduous vine thicket on coastal dunes
and basalt scree slopes (Pedley 1987; Russell-
Smith & Dunlop 1992; Liddle et al. 1994;
Department of Planning & Infrastructure
2010). It is also found in the Philippines,
Indonesia (Java) and Timor-Leste (Hyland et
al. 2003; Cowie 2006).

A survey of the vegetation of the Torres
Strait Islands, north Queensland, Australia,
was carried out in 2007 (Stanton et al.
2008). The survey’s primary objective was
to map vegetation communities at 1: 25,000
and Regional Ecosystems at 1: 50,000,
supplemented by floristic inventory and
collections of voucher specimens for
Australian herbaria. Collections of Luvunga
monophylla from lama (also known as Yam
Island) were made in October 2007 (Fell 8851
& Stanton [BRI]). This occurrence extends
its geographical range eastwards from the
eastern Arnhem Land coast of the Northern
Territory.

lama (Yam Island)

lama is a continental island of 186 ha located
at 9°54'S 142°46'E. It is situated 91 km north-
north-east of Cape York Peninsula and belongs
to the Central Group of Torres Strait Islands.
The island is the homeland of the Iamalgal
people. As at the 2006 Census, the population
was 309, of whom over 90% were Indigenous
(Australian Bureau of Statistics 2007). Land
tenure is Deed of Grant in Trust (DOGIT) and
Native Title.

Accepted for publication 8 August 2011

432

The primary geology of the island is granite
with a range of Holocene aged landforms
fringing its margins. Coarse-grained granite,
which forms the dominant landform feature,
represents the most easterly extension of the
Badu Granite batholith. Soil development is
typically skeletal, with large granite boulders
on lower slopes and littoral margins. Small
areas of acid volcanic rock form a headland
in the island’s east. Holocene features include
ridges of coralline sand and fine-grained
estuarine deposits associated with a broad
embayment on the island’s northern coast
(Willmott & Powell 1977; Stanton et al.
2008).

Remnant vegetation occurs over 75%
of the island with the balance supporting
regrowth, exotics, bamboo groves and cleared
land. Broad vegetation types are deciduous
and semi-deciduous vine forest and thicket,
Acacia dominated open forests and woodlands,
shrublands and shrubland complexes, coastal
dune complexes, grasslands and mangrove
forest (Stanton et al. 2008).

Luvunga monophylla on lama

Our vegetation survey recorded Luvunga
monophylla at eight sites on the island,
within open forest dominated by Acacia
auriculiformis A.Cunn. ex Benth. and in
deciduous vine forest and semi-deciduous
vine thicket. These vegetation types occur
on rocky granitic hillslopes and the Acacia
dominant vegetation is considered to have
developed in response to past disturbance.

The Acacia open forest habitat is endemic
to lama and the Torres Strait Islands and is
mapped as Regional Ecosystem 3.12.35f
(Stanton et al. 2008). Vine forest species
such as Canarium australianum F.Muell.,
Terminalia subacroptera Domin, Bombax
ceiba var. leiocarpum A.Robyns and
Diospyros hebecarpa A.Cunn. ex Benth.
are typical canopy associates. Groves of
the naturalised Bamboo ( Bambusa vulgaris
Schrad.) and scattered Mango trees ( Mangifera
indica L.) occur throughout this habitat and
are indicative of a long history of traditional
use by local people.

Austrobaileya 8(3): 431-434 (2011)

Deciduous vine forests and semi-deciduous
vine thickets (RE3.12.35a and RE3.12.21a)
are extensive on the lower exposed rocky
slopes and feature canopy species such as
Erythrina insularis F.M.Bailey, Antiaris
toxicaria var. macrophylla (R.Br.) Corner,
Terminalia subacroptera , Canarium
australianum , Bombax ceiba var. leiocarpum ,
Acacia auriculiformis , Manilkara kauki (L.)
Dubard, Gyrocarpus americanus Jacq. subsp.
americanus, Diospyros sp. (Mt White PI.
Forster PIF14415), Sterculia quadrifida R.Br.
and Premna dallachyana Benth.

The habitats in which Luvunga monophylla
occurs are in good condition and at the time of
survey were relatively undisturbed and free of
weeds. A conservative estimate of population
size for Luvunga monophylla is between 50
and 100 individuals. The area of extent is 50
ha.

Discussion

The nature of the disturbance history in the
Acacia open forest is unknown, although the
community would be expected to burn in hot
conditions. Old charred stumps indicate that
it is occasionally burnt, largely in a wildfire
regime. Resilience to fire is documented
from Northern Territory populations of
Luvunga which have an ability to resprout
from epicormic and basal lignotubers and
display less than 30% mortality rate when
subject to 100% leaf scorch. Life span is over
20 years with first seeding occurring between
6-20 years (Tropical Savannas Cooperative
Research Centre 2010).

The fact that the Acacia dominant
community is found only on lama presents
another puzzle. The nearby continental islands
of Mabuyag (Mabuiag Island) and Gebar
are dominated by open and closed forests of
Welchiodendron longivalve (F.Muell.) Peter
G. Wilson & J.T.Waterh., although not a single
individual of this species was found on lama.
This may suggest that forests on lama have
developed independently from forest types
on these neighbouring islands (Stanton et
al. 2008). Extensive field surveys by Stanton
et al. (2008) in similar habitats across the
majority of the Torres Strait islands did not
locate Luvunga elsewhere.

Fell & Stanton, Luvunga monophylla , new for Queensland

Prior to European contact the people
of lama practised traditional horticulture
growing crops such as banana, yam and sweet
potato (Haddon 1935; Neal 1989; Fuary 1991)
with archaeological evidence suggesting that
some gardening took place on rocky hillslopes
(Neal 1989). It is likely that a combination of
human disturbance, wind and fire have played
a role in the development of Luvunga habitat.

No uses are documented for Australian
populations of Luvunga monophylla.
Specimens were shown to numerous members
of the community during our visit to lama in
November 2010. Although they recognized the
plant, they stated that they had no name or use
for it. In Java, it is occasionally cultivated as a
hedge plant (Kruse 2001: 1013). In India, the
dried fruits of Luvunga scandens Roxb. are
used in the production of a medicinal oil said
to be effective in treating dermal infections
and baldness (Garg & Jain 1999) and the roots
and fruits are used for treatment of scorpion
stings (Lien et al. 2002).

It is interesting to note that the occurrence
of Luvunga monophylla across northern
Australia appears to be coterminous with
those areas visited in the past by Macassan
beche-de-mer fishermen from southern
Sulawesi in what is now Indonesia, Timor-
Leste and between Cape Leveque in Western
Australia and the Sir Edward Pel lew Group
in the Northern Territory (Macknight 1969: 2,
27; Stacey 2007: 58). There is no firm evidence
that this trade included Torres Strait, although
there is an 1881 record of an Ambonese prau
(boat) being driven into the region during
the northwest monsoon season (Anonymous
1881). This raises the possibility that Luvunga
monophylla was introduced to Australia by
human vectors.

Conservation status of Luvunga
monophylla

In the Northern Territory this species is
assigned ‘least concern’ status under the
Territory Parks and Wildlife Conservation
Act 2000, and in Western Australia it is
‘Not Threatened’ according to the Wildlife
Conservation Act 1950.

433

Cowie (2006) records the taxon from
dry deciduous forest in Timor-Leste,
noting that the forest type is under ongoing
pressure from conversion to agriculture.
The heavy exploitation of dry deciduous
forests elsewhere in the Timor region
for swidden agriculture has resulted in
large-scale conversion to anthropogenic
grassland -Chromolaena shrubland (Cowie
2006). The extent of its occurrence and
condition in habitats in the Philippines and
Java is not known.

Potential threats to the lama population
include habitat disturbance from infrastructure
and residential development, weed incursion
and perhaps wildfire. Pressures on the limited
availability of land suitable for housing and
infrastructure development is intensified
by population growth and coastal erosion
associated with regular tidal surges. This
presents a considerable challenge for the
island community and planning agencies.

In combination, these threats represent a
risk of stochastic extinction of the species in
Queensland due to its small population size
and highly localised occurrence. Accordingly,
the species should be assigned regional
significance in the Cape York Peninsula
Bioregion. Integration of ecological and
genetic studies will be required to determine
the management requirements of Luvunga
monophylla on lama. Further assessments of
population size, phenology and recruitment is
achievable at the local level with the assistance
of the local ranger program.

Acknowledgements

The field survey on lama was funded by the
Land and Sea Management Unit of the Torres
Strait Regional Authority. Access to lama was
gratefully provided by the Magani Lagaugal
(Torres Strait Islanders) Corporation. Advice
on identification and species distribution
was provided by the staff of the Queensland
Herbarium. Comments on an earlier version
of this paper from Garrick Hitchcock and Rob
Neal are greatly appreciated.

434

References

Anonymous (1881). The Brisbane Courier 21 February:
3.

Australian Bureau Of Statistics (2007). Population
Distribution, Aboriginal and Torres Strait
Islander Australians, Australia - 2006. Cat.
No. 4705.0. Australian Bureau of Statistics:
Canberra.

Cowie, I. (2006). A Survey of Flora and Vegetation of
the Proposed Jaco-Tutuala-Lore National
Park, Timor-Leste (East Timor). Report to
Birdlife International from NT Herbarium
(DNA) Department of Natural Resources,
Environment and the Arts: Palmerston, N.T.
Version 1.0. Available from: http://www.dpif.
nt.gov.au/nreta/publications/wildlife/science/

pdf/2006CowieI.pdf

Department Of Environment And Conservation,
Western Australian Herbarium (2010).
Florabase. Online at http://florabase.calm,
wa.gov.au/ Accessed 30 October 2010.

Department Of Planning And Infrastructure (2010).
Australian Virtual Herbarium. NT Herbarium
map tool. Northern Territory DPI, Darwin.
Available from: http:/ipe.nt.gov.au/cgi-bin/avh.
cgi.

Garg, S.C. & Jain, R. (1999). Antifungal activity of
Luvunga scandens against some keratinophilic
fungi. Indian Journal of Pharmaceutical
Sciences 61: 248-249.

Fuary, M.M. (1991). In so many words: an ethnography
of life and identity on Yam Island, Torres Strait.
PhD thesis, James Cook University of North
Queensland: Townsville.

Haddon, A.C. (1935). Reports of the Cambridge
anthropological expedition to Torres Straits.
Vol. 1. general ethnography. Cambridge
University Press: Cambridge.

Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray,
B. & Elick, R.W. (2003). Australian tropical
rain forest plants: trees and shr ubs and vines.
CSIRO Publishing: Melbourne.

Kruse, J. (2001). Rutaceae. In P. Hanelt (ed.), Mansfeld’s
encyclopedia of agricultural and horticultural
crops, pp. 996-1038. Springer Verlag: Berlin.

Kubitzki, K., Kallunki, J.A., Duretto, M. & Wilson,
PG. (2010). Rutaceae. In K. Kubitzki (ed.).
The Families and Genera of Vascular Plants.
X. Flowering Plants Eudicots Sapindales ,
Cucurbitales , Myrtaceae , pp. 276-356. Springer
Verlag: Berlin/Heidelberg.

Austrobaileya 8(3): 431-434 (2011)

Liddle, D.T., Russell-Smith, J., Brock, J., Leach, G.J.
& Connors, G.T. (1994). Atlas of the vascular
rainforest plants of the Northern Territory.
Flora of Australia Supplementary Series No.
3. Australian Biological Resources Study:
Canberra.

Lien, T.P, Kamperdick, C., Schmidt, J., Adam, G. & Van
Sung, T. (2002). Apotirucallane triterpenoids
from Luvunga sarmentosa (Rutaceae)
Phytochemistry 60: 747-754.

Mabberley, D.J. (1998). Australian Citreae with notes
on other Aurantioideae (Rutaceae). Telopea 7:
333-344.

Macknight, C.C. (1976). The voyage to Marege’:
Macassan Trepangers in Northern Australia.
Melbourne University Press: Carlton.

Neal, R.A. (1989). An Archeological Inspection of
Alternative Telecom Locations on Mabuiag
and Yam Islands, Torres Strait. Unpublished
report to Department of Community Services
and Ethnic Affairs. Pirriport Pty Ltd: Helidon,
Queensland.

Pedley, L. (1987). Paramignya Wight (Rutaceae:
Citreae) in Australia. Austrobaileya 2: 416.

Russell-Smith, J. & Dunlop, C.R. (1992). The status of
monsoon vine forests in the Northern Territory:
a perspective. In G.L. Werren & A.P Kershaw
(eds.) The rainforest legacy. Australian
National Rainforests Study. Vol. 1, The Nature,
Distribution and Status of Rainforest types, pp.
227-288. Australian Government Publishing
Service: Canberra.

Stacey, N. (2007). Boats to burn: Bajo fishing activity
in the Australian fishing zone. Asia-Pacific
Environment Monograph 2. ANU E Press:
Canberra.

Stanton D.J., Fell, D.G. & Gooding, D.O. (2008).
Vegetation Communities and Regional
Ecosystems of the Torres Strait Islands,
Queensland, Australia. Unpublished report
to the Torres Strait regional Authority, Land
and Sea Management Unit. 3D Environmental
Trust: Brisbane.

Tropical Savannas Cooperative Research Centre
(2010). North Australian Land Manager
Website. Available from: http://landmanager,
org.au/view/311424/fire - responses-of--

luvunga-monophvlla.html

Willmott, W.F. & Powell, B.S. (1977). Torres Strait-
Boigu-Daru, Queensland 1:250 000 Geological
Series - Explanatory Notes, Sheets SC/ 54-12,
SC/ 54-7 and SC/ 54-8. Bureau of Mineral
Resources, Geology and Geophysics. Australian
Government Publishing Service: Canberra.

Austrobaileya 8(3): 435-437 (2011)

435

SHORT COMMUNICATION

Hydrocharis dubia (Blume) Backer (Hydrocharitaceae)
is an alien species in Australia

A.R. Bean

Queensland Herbarium, Department of Environment & Resource Management, Brisbane Botanic
Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia. Email: tony.bean@derm.qld.
gov.au

Hydrocharis dubia (Blume) Backer, is an
aquatic plant with a white 3-petaled flower
that occurs widely in eastern Asia. It has
been accepted as an indigenous species in
Australia but recently this has been repeatedly
questioned by weed control authorities. This
note examines the evidence relating to its
origin status and concludes that it is not native
in Australia.

The genus Hydrocharis L. was revised by
Cook & Luond (1982), with the recognition of
three species,//. morsus-ranaeL.,H. chevalieri
(De Wild.) Dandy and H. dubia. Hydrocharis
morsus-ranae is widely distributed in Europe,
with scattered occurrences in western Asia. It
has become a serious weed in south-eastern
Canada (Cook & Luond 1982) and in the
north-eastern U.S.A. (Catling et al. 2003;
O’Neill 2007). Hydrocharis chevalieri is
restricted to central Africa, and is not recorded
as a weed. Hydrocharis dubia is widely
distributed in south-eastern Asia, from India
to Japan and south to Java and New Guinea,
and in Australia (Cook & Luond 1982). It has
been referred to as a “noxious weed” in Japan
(Oki 1994). Shaffer-Fehre (1991) transferred
Hydrocharis dubia to Limnobium, a genus
which until then had comprised two species
confined to the Americas; however, this
transfer has not been followed in the Flora
of Australia (Jacobs & McColl 2011) and is
not supported by molecular data (Les et al.
2006).

Hydrocharis dubia was first recorded
for Australia by Bentham (1873) under the
misapplied name// morsus-ranae. The record

Accepted for publication 8 September 2011

was based on a specimen (now at K) collected
by J. Bidwill, from “Wide Bay” between 1848
and 1853. This specimen was cited by Cook
& Luond (1982) under Hydrocharis dubia.
Bentham (1873) entertained the idea that it
could be an introduced species in Australia.

Bailey (1902) recorded Hydrocharis
morsus-ranae from “the still waters about the
Brisbane River”. There is a specimen record
of Hydrocharis dubia on the BRI database
with the locality “Brisbane River”, but the
specimen cannot currently be located.

Hydrocharis dubia has been found at 13
Australian locations in Queensland (Qld) and
New South Wales (NSW) based on herbarium
records (Table 1).

Hydrocharis dubia (as Limnobium dubium
by some authors) has been regarded as an
indigenous species in Australia (Aston 1973;
Stanley & Ross 1989; Jacobs 1993; Holland
& Hansen 2009; Jacobs & McColl 2011). The
reasons for this have not been expounded, but
they are probably two-fold: that the species
is indigenous to south-east Asia, and natural
dispersal from that region to Australia is not an
unreasonable hypothesis; and that the species
is included in Bentham’s Flora Australiensis.
Thomas & McDonald (1989) gave
Hydrocharis dubia a status of ‘2V’, meaning
a Vulnerable species with a geographic range
in Australia of less than 100 km. Presumably
the authors, at that time, considered it native
to south-eastern Queensland. Subsequently
it was classified as “Vulnerable” under the
Queensland Nature Conservation Act 1992,
and “Vulnerable” under the Environment
Protection and Biodiversity Conservation Act
1999.

436

Austrobaileya 8(3): 435^37 (2011)

Table 1. Locations for Hydrocharis dubia in Australia over time

Localities

Time period of collections

Wide Bay, Qld

1848-1853

Clarence River, NSW

1882

Brisbane River, Qld

pre-1902

Harrisville, Qld

1960, 1985, 1986

Bountiful Creek, Richmond River, NSW

1967

Frederickton near Kempsey, NSW

1968

Beaudesert, Qld

1971, 1977

Toomba, Qld

1977, 1996, 1997, 2001, 2008

16 km W of Casino, NSW

1978

Ayr, Qld

1994

Rosewood, Qld

1999, 2001, 2009

Fletcher Ck, Qld

2002, 2004

Giru, Qld

2004

The following observations are made on
Hydrocharis dubia in Australia drawing on
criteria used by Bean (2007) to assess its
origin status:

1. it grows in disturbed or modified sites, and
has not been found in remote “intact” areas.
This is consistent with a species introduced
by man, and transported to other sites via
machinery or with soil, or dumping of plants

2. it is persistently invasive (one label reports
it covering two dams), and is not in ecological
balance with surrounding biota

3. it does not appear to have any pests or
diseases, suggesting it has arrived recently in
evolutionary terms

4. it has substantially expanded its
geographical range over the last 40 years (see
list of locations above)

5. its distribution is highly discontinuous, and
the discontinuities cannot be explained by
climatic or other natural criteria

6. the initial locations in Australia are highly
disjunct geographically from nearest known
occurrences in Asia

7. the first record for Australia (Wide Bay,
between 1848 and 1853, by JohnBidwill) is not
early enough to preclude human introduction

Hydrocharis dubia fails all of the
ecological criteria for an indigenous species,
and the historical criteria are equivocal. In the
key of Bean (2007) it readily keys to “alien”
(lx, 2).

In conclusion, there is a strong weight
of evidence to support an alien status
for Hydrocharis dubia with regard to its
occurrences in Australia. I recommend that
this species be recorded as naturalised for
Australia, and de-listed from the Queensland
Nature Conservation Act 1992 , and the
Environment Protection and Biodiversity
Conservation Act 1999.

References

Aston, H.I. (1973). Aquatic Plants of Australia.
Melbourne University Press: Carlton.

Bailey, F.M. (1902). Hydrocharis. InFlora of Queensland
5: 1510. H. J. Diddams & Co.: Brisbane.

Bean, A.R. (2007). A new system for determining
which plant species are indigenous in Australia.
Australian Systematic Botany 20: 1-43.

Bentham, G. (1873). Hydrocharis. In Flora Australiensis
6: 256. L. Reeve & Co.: London.

Catling, P.M., Mitrow, G., Haber, E., Posluszny, U.
& Charlton, W.A. (2003). The biology of
Canadian weeds. 124. Hydrocharis morus-
ranae L. Canadian Journal of Plant Science 83:
1001-1016.

437

Bean, Hydrocharis dubia alien in Australia

Cook, C.D.K. & Luond, R. (1982). A revision of the
genus Hydrocharis (Hydrocharitaceae). Aquatic
Botany 14: 177-204.

Holland, A.E. & Hansen, D. (2009). Species
Information - Hydrocharis dubia. Wetland
Info, Queensland Government. http://www.
epa. qld .gov. au/wetlandinfo/site/factsfigures/
FloraAndFauna/Flora/IndicatorSpeciesList/121
14.html Accessed 29 March 2011.

Jacobs, S.W. (1993). Hydrocharitaceae. In G. Harden
(ed.). Flora of New South Wales 4: 11-16. New
South Wales University Press: Sydney.

Jacobs, S.W. & McColl, K.A. (2011). Hydrocharis. In
Flora of Australia 39: 21-22. ABRS/CSIRO:
Melbourne.

Les, D.H., Moody, M.L. & Soros, C.L. (2006). A
reappraisal of phylogenetic relationships in the
monocotyledon family Hydrocharitaceae. Aliso
22: 211-230.

Oki, Y. (1994). Integrated Management of aquatic
weeds in Japan. Food & Fertilizer Technology
C enter, http: II www. agnet. org/1 ibrary/bc/45008/
Accessed 29 March 2011.

O’Neill, C.R. (2007). European Frog-bit ( Hydrocharis
morsus-ranae ) - Floating invader of Great
Lakes Basin Waters. NYSG Invasive Species
Factsheet Series: 07-1. http://www.seagrant.
sunysb.edu/ais/pdfs/Frog-bitFactsheet.pdf
Accessed 29 March 2011.

Shaffer-Fehre, M. (1991). The endotegmen tuberculae:
an account of little-known structures from
the seed coat of the Hydrocharitoideae
(Hydrocharitaceae) and Najas (Najadaceae).
Botanical Journal of the Linnean Society 107:
169-188.

Stanley, T.D. & Ross, E.M. (1989). Flora of south¬
eastern Queensland , Volume 3. Department of
Primary Industries: Brisbane.

Thomas, M.B. & McDonald, W.J.F. (1989). Rare and
threatened plants of Queensland , 2 nd edition.
Department of Primary Industries: Brisbane.

Austrobaileya 8(3): 438-440 (2011)

438

SHORT COMMUNICATION

Reinstatement of Vigna suberecta Benth. (Fabaceae: Phaseoleae)

Ailsa E. Holland

Queensland Herbarium, Department of Environment & Resource Management, Brisbane Botanic
Gardens, Mt Coot-tha, Toowong, Queensland 4066, Australia. Email: Ailsa.Holland@derm.qld.gov.au.

The genus Vigna Savi is a pantropical,
herbaceous legume genus of about 200
species distributed throughout the tropics
and subtropics of both hemispheres. In
Australia, there are a number of endemic
Vigna taxa, including those in the wide
ranging V lanceolata Benth. complex (Lawn
& Holland 2003).

Vigna lanceolata and Vigna suberecta
Benth. were both originally described by
Bentham in Mitchell (1848). On the 10 th of
October 1846 Mitchell arrived at the summit
of Mt Farraday (Mitchell 1848: 347) where
he first encountered “the yellow Vigna
lanceolata ” and collected specimen number
376, the left hand specimen on Kew sheet
H/1215/89 4, labelled “ Vigna lanceolata
Benth. in Mitch Austr. Trop. p. 350. 1846. 376.
Oct 10 402, small twining herb, fl yellow, top
of Mount Farraday” and annotated as the
type by Marechal, although he later refers
to specimen 370 (Marechal et al:. 1978: 168)
which is probably an error.

On 13 th of October 1846, Mitchell had
advanced to Mt Owen (Mitchell 1848: 350),
where he found “a new species of Vigna
with yellow flowers” and collected specimen
number 401. This is the right hand specimen
labeled as “Oct 13 420 on Mt Owen... probably
label changed with Swainsona phasoides ”,
on another Kew sheet H/1215/89 3 and is the
same taxon as the previous. This species was
described by Bentham in the footnote on this
page (translated): “V. LANCEOLATA (Benth.
MS.). Glabrous, twining. Leaflets lanceolate,
reticulate, entire, hastate or lobed at the base.
Peduncles much longer than leaves, with few
flowers clustered at apex. Calyx glabrous,
campanulate, the lobes shorter than the tube;
keel beak acute”

Accepted for publication 9 September 2011

Mitchell arrived on the Moonie River
on the 18 th November 1846 (Mitchell 1848:
387) and collected another specimen “in the
thick forest, a new species of Vigna very
near V. lanceolata , though very different in
habit”. This is specimen number 466, Kew
sheet H/1215/89 5, label “1846. 466. Vigna
suberecta Benth. in Mitch Austral, trop.”
This species was described by Bentham in
a footnote (Mitchell 1848: 388) translated
as: “V. SUBERECTA (Benth. MS.); slightly
pubescent, sub-erect, branched. Leaflets
broadly lanceolate leaflets, entire, hastate or
3-lobed at base. Peduncles somewhat shorter
than the leaves, few-flowered at the apex.
Calyx pubescent, campanulate; calyx lobes
subequal to the tube; keel beaked, acute. Pod
sparsely pubescent .”

Bentham later (1864) placed Vigna
suberecta in synonymy under V. lanceolata ,
but did not give a reason. Examination
of the type material (the sheets indicated
above) shows that the specimens labelled “V
lanceolata ” (376 and 401) do indeed belong
to the same taxon, but the specimen labelled
“E suberecta ” (466) is quite distinct and so
this name is here reinstated for a relatively
widespread species.

Vigna suberecta Benth., in T.L.Mitchell,
J. Exped. Trop. Australia[Mitchell] 388 (1848).
Type: subtropical New Holland [Queensland.
Darling Downs District: Moonie River], [18
November] 1846, T.L. Mitchell 466 (holo: K).

Vigna sp. (Jimbour A.R.Bean 12534); Holland
& Pedley (2007: 84).

Herb with perennial tuberous rootstock; tubers
cylindrical, up to 15 mm diameter. Aerial
parts annual or short-lived perennial. Stems
to 80 cm long, initially semi-erect, becoming
twining in upper parts at maturity, with a few
appressed, retrorse white hairs to 0.8 mm

Holland, Reinstatement of Vigna suberecta (Fabaceae)

long. First leaves unifoliolate, petiolate; blade
narrowly triangular, 2-2.5 cm long, 4-7 mm
wide, truncate to slightly sagitate at base.
Remaining leaves trifoliolate; petiole 2.2-
5.7 cm long; rachis 5-15 mm long; stipels
lanceolate, 1-2.3 mm long. Terminal leaflets
lanceolate, becoming linear towards the plant
apex, 2.5-7 cm long, 3-17 mm wide; 3 to 13
times as long as wide, apex acute, base cuneate
with rounded or acute lobes, 1-4 mm long
and wide; upper surface glabrous or nearly
so, lower surface very sparsely appressed
hairy; midrib uniformly coloured (not silver
striped), secondary nerves forming an angle
of 45-80° to the midrib, looped at margin.
Lateral leaflets similar, slightly shorter
than terminal leaflet. Stipules triangular
to lanceolate, 1.8-3.5 mm long, 3 or 5-
veined, acute, base ± cordate. Subterranean
inflorescences absent. Aerial inflorescences
with 2-10 flowers crowded towards the apex;
flowers developing sequentially in pairs with
a swollen extrafloral nectary between each
pair. Peduncle terete, striate, 2-16 cm long;
pedicel 1-2.2 mm long; bracts and bracteoles
ovate, acute, 0.8-1.2 mm long; bracteoles
inserted at base of calyx. Calyx campanulate,
2.2-4.5 mm long, glabrous or minutely ciliate;
tube 1.5-2.5 mm long; lobes triangular, equal
to or slightly shorter than tube, upper two
joined almost to apex, lower one longest.
Corolla pale yellow, glabrous. Standard
transversely elliptic, emarginate, 6-9 mm
long, 10—14 mm wide; the inner surface with
two U-shaped pockets with vertical ridges, c.
2 mm long; apex emarginate; base somewhat
auriculate; claw 1-1.5 mm long. Wings 6-10
mm long, 4-7 mm wide, expanding from
a narrow base into a broadly rounded apex
covering the keel; upper margin with a small
spur near base; lower margin without a spur;
claw 1.5-2 mm long. Keel 5-9 mm long, 4-6
mm wide; upper margin with a shallow pocket
near base; lower margin strongly curved;
beak strongly upturned and twisted through
c. 180°; claw 2-3 mm long. Filament tube
c. 10 mm long, strongly upturned; filaments
free for 1/3 - 1/2 their length; anthers 2-3
mm long. Ovary pubescent, 4-6 mm long;
style terete, twisted, anterior face hairy for
1/3—1/2 the length; stigma situated near apex,
the style slightly prolonged past stigma into a

439

beak 2-3 mm long, straight or slightly curved
by less than 90° Pods slightly curved, 2.5-4
cm long, 3-6 mm wide, sparsely puberulent
when young, glabrescent, brown or black at
maturity. Seeds 6-10, oblong, dark brown or
reddish, 3-4.2 mm long, 3-3.7 mm wide, 2-
2.6 mm thick; hilum central, 1-1.3 mm long.

Additional selected specimens examined : Queensland.
Leichhardt District: 169 km from Clermont toward
Mackay on Peak Downs Highway, Dec 1987, Lawn &
Cottrell ACC906 (BRI); 9 km N of Capella, Mar 1995,
Fensham 2769 (BRI); 9 miles [14.4 km] SSW of Comet,
Feb 1960, Johnson 1382 (BRI). Burnett District:
Wooroolin, Portion 31, Nov 1961, Bailey s.n. (BRI
[AQ239116]). Maranoa District: Roma, Mar 1936, Blake
10854A (BRI). Darling Downs District: Marnhull, SE of
Jandowae, Dec 2001, Bean 18199 (BRI); Jimbour Plain,
Mar 1951, BissetS758 (BRI); CSIRO experimental area,
Nandi, Dalby, Apr 1983, Watkinson ACC207 (BRI);
19 km SSW of Dalby, Feb 1995, Fensham 1909 (BRI);
Jondaryan, Feb 1935, Blake 7753 (BRI); 8 km S of Cecil
Plains turnoff (E of Dalby) at Duncan’s road turnoff,
Nov 2001, Holland 1388 et al. (BRI, US); Cecil Plains to
Milmerran Road at River Road turnoff. Mar 1987, Lawn
et al. ACC261 (BRI). New South Wales. Yalleroi, s.dat.,
Wilson ACC210 (BRI); Moree, Mar 1962, Cutting s.n.
(NSW); Delungra, Feb 1968, Gilroy s.n. (NSW); Wee
Waa, c. 32 km NW of Narrabri, Feb 1981, Nehrkorn s.n.
(NSW); Edgeroi, 16 miles [25.6 km] N of Narrabri, Feb
1966, Quick s.n. (NSW); Gunnedah, May 1938, O’Reilly
s.n. (NSW).

Distribution and habitat: Vigna suberecta
is a widespread species occurring mainly in
the Brigalow Belt bioregion of Queensland
from Clermont in the north to as far west as
Roma, and as far south as Gunnedah in New
South Wales. It is confined to heavy clay soils
in grassland or open eucalypt woodland with
grassy under storey.

Phenology: Flowers in summer, mostly
between November and March and fruits
through to June, depending on rainfall.

Notes: Vigna suberecta is distinguished
from V. lanceolata by the leaflets lacking a
silver midrib, wing petals rounded, keel beak
twisted c. 180°, and seeds with a longer (1-1.3
mm) hilum (Table 1).

While Vigna suberecta is a semi-erect
herb in the early stages, it becomes vine-like
and twining at maturity. Vigna suberecta
often resprouts from tubers in spring, or after
fire.

440 Austrobaileya 8(3): 438-440 (2011)

Table 1. Morphological character differences between Vigna lanceolata Benth. and
V. suberecta Benth.

Character

V lanceolata

V suberecta

Underground inflorescences

yes

no

leaflets midrib silver striped

yes

no

standard with raised vertical ridges

no

yes

wing shape

triangular

rounded

keel beak twist

c. 90°

c. 180°

seed hilum length

0.7-1.1 mm

1-1.3 mm

Acknowledgements

I would like to thank the director, Royal
Botanic Gardens Kew for the loan of type
material, and the director of the National
Herbarium of New South Wales for the
specimen loan. Thanks to Prof R. Lawn and
Dr L. Bielig (both JCU) for their support
and encouragement, and to Peter Bostock
for the excellent Latin translation program
TRANSLAT. This work was done in partial
fulfillment of a PhD degree at James Cook
University (JCU).

References

Bentham, G. (1848). In T.L.Mitchell, Journal of
an Expedition into the Interior of Tropical
Australia in Search of a Route from Sydney
to the Gulf of Carpentaria. Longman, Brown,
Green & Longmans: London.

- (1864). Vigna. In Flora Australiensis 2: 260.

London: Reeve & Co.

Holland, A.E. & Pedley, L. (2007). Fabaceae. In
P.D.Bostock & A.E.Holland (eds). Census
of the Queensland Flora 2007, pp. 72-85.
Environmental Protection Agency: Brisbane.

Lawn, R.J. & Holland, A.E. (2003). Variation in
the Vigna lanceolata complex for traits of
taxonomic, adaptive or agronomic interest.
Australian Journal of Botany 51: 295-308.

Marechal, R., Mascherpa, J.M. & Stainier, F. (1978).
Etude taxonomique d’un groupe complexe
d’especes des genres Phaseolus et Vigna
(Papilionaceae) sur la base de donnees
morphologiques et polliniques, traitees par
l’analyse informatique. Boissiera 28: 1-273.